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Microbiology

Chapter Concepts
28.1 Viruses
Viruses are noncellular. Viruses do reproduce,
but may live inside living cells. 574
Viruses are obligate intracellular parasites
because they must reproduce inside host
cells. 574

28.2 Kingdom Monera


The monera are prokaryotes of small size but are
very diverse metabolically. 577
Most bacteria are heterotrophic by absorption
but the cyanobacteria are important
photosynthesizers. 578

28.3 Kingdom Protista


The protists include eukaryotic unicellular
organisms and some related multicellular
forms. 583
Among protists, algae are the plantlike protists;
protozoans are the animal-like protists; slime
molds and water molds are the funguslike
protists. 583

28.4 Kingdom Fungi


Fungi are multicellular eukaryotes with a
lamentous body. 591
Fungi are the most complex organisms to rely on
saprotrophic nutrition. 591
Most bacteria dont cause illness, but Salmonella typhimurium
is responsible for food poisoning. Poultry, eggs, pork, beef, and oysters are all carriers of this bacterium.

573

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28-2

Evolution and Diversity

lace a single drop of pond water under a microscope


and you can see an amazing menagerie of unicellular
organisms, such as protozoans and algae. Fungi and
bacteria also abound in nature. Fungi feed on organic matter in the soil, but bacteria are even found in inorganic
environmentsin water boiling out of the oceans volcanic
vents and within rocks miles below the surface of the earth,
for example. Closer to home, microbes can live in the
crevices of your kitchen cutting board or within the food
youre eating. Set a loaf of bread, free of preservatives, on
the counter and youll observe colonies of bacteria and fungi
in no time.
Most microbes are harmless to people, and many are
benecial. Some viruses, bacteria, and fungi, however, make
a home inside the tissues or cells of people. To combat
these microbial invaders, researchers desperately hunt for
drugs, such as antiviral agents, and antibiotics, which kill
bacteria and fungi. Some antibiotics are extracted from microbes themselves, and scientists are now hopeful of reviving the decades-old idea of using bacteria-destroying
viruses against bacteria.
This chapter will introduce you to the myriad of
organisms that inhabit the microscopic world.

ample, bacteriophages infect only bacteria, the tobacco


mosaic virus infects only plants, and the rabies virus infects
only mammals. Some human viruses even specialize in a
particular tissue. Human immunodeciency viruses (HIV)
enter specic types of blood cells, the polio virus reproduces
in spinal nerve cells, the hepatitis viruses infect only liver
cells. What could cause this remarkable parasite-host cell relationship? It is now believed that viruses are derived from
the very cell they infect; the nucleic acid of viruses came
from their host cell genomes! Therefore, viruses must have
evolved after cells came into existence, and new viruses are
probably evolving even now.
Viruses, like other organisms, can mutate, and this habit
can be quite troublesome because a vaccine that is effective
today may not be effective tomorrow. Flu viruses are well
known for mutating, and this is why you have to have a u
shot every yearantibodies generated from last years shot
are not expected to be effective this year.
Viruses are nonliving particles that reproduce only
inside specic host cells.
protein unit

28.1

Viruses

Viruses are not included in the classication table found in


Appendix B because they are noncellular and should not be
classied with cellular organisms. Viruses are generally
smaller than 200 nm in diameter and therefore are comparable in size to that of a large protein macromolecule. Many
can be puried and crystallized, and the crystals can be
stored just as chemicals are stored.

DNA

capsid

Structure of Viruses

fiber

A virus always has at least two parts: an outer capsid composed of protein units, and an inner core of nucleic acid
either DNA or RNA (Fig. 28.1). The viral genome has at most
several hundred genes; a human cell contains thousands of
genes. A virus may also contain various enzymes for nucleic
acid replication. The capsid is often surrounded by an outer
membranous envelope, which is actually partially composed
of the hosts plasma membrane. The classication of viruses
is based on (1) type of nucleic acid, including whether it is
single stranded or double stranded, (2) viral size and shape,
and (3) the presence or absence of an outer envelope.

Parasitic Nature
Viruses are obligate intracellular parasites. In order to have a
ready supply of animal viruses in the laboratory, they are
sometimes injected into live chick embryos. Today, however,
it is more customary to infect cells that are maintained in tissue culture. Viruses infect all sorts of cellsfrom bacterial
cells to human cellsbut each type is very specic. For ex-

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Figure 28.1 Adenovirus.


An adenovirus is a DNA virus with a polyhedral capsid and a ber at
each corner.

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Chapter 28

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Replication of Viruses
Viruses are specic to a particular host cell because portions of the capsid (or the spikes of
the envelope) bind in a lock-and-key manner
with a receptor on the host cell plasma membrane. After viral nucleic acid enters the cell, it
takes over the metabolic machinery of the host cell
so that more viruses are produced.

Replication of Bacteriophages
Bacteriophages, or simply phages, are viruses
that parasitize bacteria; the bacterium in
Figure 28.2 could be Escherichia coli, which
lives in our intestines. In the lytic cycle, the
host cell undergoes lysis, a breaking open of
the cell to release new viruses. In the lysogenic
cycle, viral replication does not immediately
occur, but replication may take place sometime
in the future. The bacteriophage, termed
lambda, is capable of carrying out both cycles.
Lytic Cycle The lytic cycle may be divided
into five stages: attachment, penetration,
biosynthesis, maturation, and release. During
attachment, portions of the capsid combine
with a receptor on the rigid bacterial cell wall
in a lock-and-key manner. During penetration,
a viral enzyme digests away part of the cell
wall, and viral DNA is injected into the bacterial cell. Biosynthesis of viral components begins after the virus brings about inactivation of
host genes not necessary to viral replication.
The virus takes over the machinery of the cell
in order to carry out viral DNA replication and
production of multiple copies of the capsid protein subunits. During maturation, viral DNA and
capsids are assembled to produce several hundred viral particles. Lysozyme, an enzyme coded
for by a viral gene, is produced; this disrupts the
cell wall, and the release of new viruses occurs.
The bacterial cell dies as a result.

bacterial
cell wall
bacterial
chromosome

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4. Maturation:
Assembly of
viral
components.

a. Lytic Cycle

2. Penetration:
Viral DNA
enters host.

3. Biosynthesis:
Viral components
are synthesized.

b. Lysogenic Cycle

prophage

Integration:
Viral DNA passed on
when bacteria reproduce.

Figure 28.2 Lytic and lysogenic cycles.


a. In the lytic cycle, viral particles escape when the cell is lysed (broken open).
b. In the lysogenic cycle, viral DNA is integrated into host DNA. At some time in
the future, the lysogenic cycle can be followed by the lytic cycle.

Lysogenic Cycle In the lysogenic cycle, the infected bacterium does not immediately produce viruses but may do so
sometime in the future. In the meantime the phage is
latentnot actively replicating. Following attachment and
penetration, viral DNA becomes integrated into bacterial
DNA with no destruction of host DNA. While latent, the

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capsid
nucleic acid

1. Attachment:
Capsid
combines
with receptor.

During the lytic cycle, a bacteriophage takes over


the machinery of the cell so that viral replication
and release occur.

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5. Release:
New viruses
leave host cell.

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viral DNA is called a prophage. The prophage is replicated


along with the host DNA, and all subsequent cells, called
lysogenic cells, carry a copy of the prophage. Certain environmental factors, such as ultraviolet radiation, can induce
the prophage to enter the lytic stage of biosynthesis, followed by maturation and release.
During the lysogenic cycle, the phage becomes a
prophage that is integrated into the host genome.
At a later time, the phage may reenter the lytic
cycle and replicate itself.

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28-4

Evolution and Diversity

Replication of Animal Viruses


Animal viruses replicate in a manner similar to bacteriophages, but there are modications. If the virus has an envelope, its glycoprotein spikes allow the virus to adhere to
plasma membrane receptors. Then the capsid and viral
genome penetrate a host cell. Once inside, the virus is uncoated as the capsid is removed. The viral genome, either
DNA or RNA, is now free of its coverings and biosynthesis
proceeds. Another difference among enveloped viruses is
that viral release occurs by budding. As the virus buds from
the cell, it acquires an envelope partially consisting of host
plasma membrane. Certain envelope components, such as
the glycoproteins that allow the virus to enter a host cell,
were coded for by viral genes. Budding does not necessarily
result in the death of the host cell.
After animal viruses enter the host cell, uncoating
releases viral DNA or RNA, and replication occurs.
If release is by budding, the viral particle acquires
a membranous envelope.

Retroviruses Retroviruses are RNA animal viruses that


have a DNA stage (Fig. 28.3). A retrovirus contains a special
enzyme called reverse transcriptase, which carries out
RNA cDNA transcription. The DNA is called cDNA because it is a DNA copy of the viral genome. Following replication, the resulting double-stranded cDNA is integrated into
the host genome. The viral DNA remains in the host genome
and is replicated when host DNA is replicated. When and if
this DNA is transcribed, new viruses are produced by the
steps we have already cited: biosynthesis, maturation, and releasenot by destruction of the cell, but by budding.

The enzyme reverse transcriptase allows


retroviruses to produce a cDNA copy of their genes
which become integrated into the host genome.

Viral Infections
Viruses are best known for causing infectious diseases in
plants and animals, including humans. Some animal viruses

gp120
reverse transcriptase

Release:
budding gives
virus an envelope.

envelope
protease

capsid
integrase

8 Maturation:
assembly of
viral components.

protease
HIV provirus

integrase

HIV particle

proteins

host DNA

viral RNA
reverse transcriptase

cDNA
6

3
4

1 Attachment: gp120
binds to CD4 receptor
and then co-receptor.

viral RNA

7 Biosynthesis:
viral components
are synthesized.

double-stranded cDNA
6 Transcription:
produces many
strands of mRNA.

2 Penetration: viral RNA


and viral enzymes enter
the host cell.
3 Reverse transcription:
produces cDNA strand.

4 Replication: produces
double-stranded cDNA.

5 Integration: viral DNA passed on


when cell reproduces.

Figure 28.3 Reproduction of the retrovirus HIV-1.


HIV-1 utilizes reverse transcription to produce cDNA (DNA copy of RNA genes). cDNA integrates into the cells chromosomes before it
reproduces and buds from the cell.

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28-5

Chapter 28

Microbiology

577

plasmid
DNA strand
in nucleoid
flagellum

ribosome
cytosol

Figure 28.4 Typical bacterial cell.

capsule

The plasma membrane of a bacterium is usually surrounded by


a cell wall and often a capsule. Note there are no organelles
except ribosomes.

cell wall
plasma membrane

a.

are specic to human cells. Of special concern are those such


as the papillomavirus, the herpes viruses, the hepatitis
viruses, and the adenoviruses, which can cause specic
types of cancer. Retroviruses are of interest because human
immunodeciency viruses (HIV), which cause AIDS, are
retroviruses. Retroviruses also cause certain forms of cancer.
At least a thousand different viruses cause diseases in
plants. About a dozen crop diseases have been attributed
not to viruses but to viroids, which are naked strands of
RNA not covered by a capsid. Like viruses, though, viroids
direct the cell to produce more viroids.
Some diseases in humans have been attributed to prions, which are protein particles that possibly can convert
other proteins in the cell to become prions. It seems that prions may have a misshapen tertiary structure and can cause
other similar proteins to convert to this shape also. Prions
are believe to cause Creutzfeldt-Jakob disease (CJD) a mental disorder that over a decade can lead to loss of vision and
speech before paralysis and death occur. Recently, prions
have been linked to a serious outbreak in Great Britain of
bovine spongiform encephalopathy (BSE), better known as
mad cow disease. Cattle feed had been supplemented with
the remains of sheep that had died of scrapie, another prion
disease. In an effort to prevent an outbreak of BSE in this
country, the importation of cattle feed, meat products, and
live cattle from Great Britain and any other BSE-affected
country has been banned since 1989.
Viruses, viroids, and prions are all known to cause
diseases in humans.

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Kingdom Monera

Kingdom Monera contains the bacteria (Fig. 28.4), which are


prokaryotes. Prokaryotic cells do not have a nucleus as do
eukaryotic cells.

fimbriae

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28.2

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Structure of Bacteria
Prokaryotic cells are very small (110 m in length and
0.71.5 m in width), and except for ribosomes, they do not
have the cytoplasmic organelles found in eukaryotic cells.
They do have a chromosome, but it is contained within a nucleoid, which has no nuclear envelope; therefore, bacteria
are said to lack a nucleus. Many bacteria have extrachromosomal rings of DNA called plasmids, which are often extracted and used as vectors to carry foreign DNA into other
bacteria during genetic engineering.
Bacteria have a cell wall containing unique amino sugars cross-linked by peptide chains. The cell wall may be surrounded by a capsule. Some bacteria move by means of
agella, and some adhere to surfaces by means of short, ne,
hairlike appendages called mbriae.

Prokaryotic cells lack a nucleus and most of the


other organelles found in eukaryotic cells.

Metabolism of Bacteria
Some bacteria are obligate anaerobes, unable to grow in the
presence of oxygen. A few serious illnesses, such as botulism and tetanus, are caused by anaerobic bacteria. Some
other bacteria, called facultative anaerobes, are able to grow
in either the presence or the absence of oxygen. Most
bacteria, however, are aerobic and, like animals and plants,
require a constant supply of oxygen to carry out cellular
respiration.

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Evolution and Diversity

a. Bacilli in pairs

b. Cocci in chains

c. A spirillum with flagella

Figure 28.5 Diversity of bacteria.


a. Bacillus (rod-shaped) bacterium. b. Coccus (round) bacterium. c. Spirillum (spiral-shaped) bacterium.

Every type of nutrition is found among bacteria except


heterotrophism by ingestion. Some bacteria are autotrophic
by photosynthesis; they use light as a source of energy to produce their own food. Cyanobacteria (see Fig. 28.6), which
photosynthesize in the same manner as plants, also give off
oxygen. Some bacteria are autotrophic by chemosynthesis;
they oxidize inorganic compounds to obtain the necessary
energy to produce their own food. Such bacteria are very important to the cycling of nitrogen in ecosystems.
Most types of bacteria are heterotrophic by absorption.
They are saprotrophs, organisms that carry on external digestion of organic matter and absorb the resulting nutrients
across the plasma membrane. In ecosystems, bacteria are
called decomposers because they break down organic matter and make inorganic nutrients available to photosynthesizers. The metabolic capabilities of bacteria are utilized for
the digestion of sewage and oil and also for the production
of such products as alcohol, vitamins, and even antibiotics.
By means of genetic engineering, bacteria are now used to
produce useful substances, such as human insulin and
growth hormone.
Bacteria are often symbiotic; they live in association
with other organisms. The nitrogen-xing bacteria in the
nodules of legumes are mutualistic, as are the bacteria that
live within our own intestinal tract. We provide a home for
the bacteria, and they provide us with certain vitamins.
Commensalistic bacteria reside on our skin, where they usually cause no problems. Parasitic bacteria are responsible for
a wide variety of plant and animal diseases.

The majority of bacteria are heterotrophic by


absorption (saprotrophic decomposers) and
contribute signicantly to recycling matter through
ecosystems.

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Classication of Bacteria
Bacteria are found in three basic shapes. (Fig. 28.5): rod
(bacillus, pl., bacilli); round or spherical (coccus, pl., cocci);
and spiral or helical-shaped (spirillum, pl., spirilli). These
three basic shapes may occur in particular arrangements. For
example, cocci may form clusters (staphylococci, diplococci)
or chains (streptococci). Rod-shaped bacteria may appear as
very short rods (coccobacilli) or as very long laments.
Traditionally bacteria are classied as either Gram-positive
or Gram-negative. Gram-positive bacteria retain a dye-iodine
complex and appear purple under the light microscope, while
Gram-negative bacteria do not retain the complex and appear
pink. Gram-positive bacteria have a thick layer of peptidoglycan on their cell wall, whereas Gram-negative bacteria have
only a thin layer. For the past 75 years bacterial taxonomy has
been compiled in Bergeys Manual of Determinative Bacteriology.
The most recent edition of Bergeys Manual divided the bacteria
into 10 major groups, which were further subdivided into order, families, genera, and species. The names of the groups
for example, Nonmotile, Gram-negative, curved bacteria or
Nonsporeforming Gram-positive rodsreect the phenotypic way the manual groups the bacteria.
A new way of classifying bacteria on the basis of rRNA
(ribosomal RNA) sequences was introduced in the 1980s.
Bacteria that share the same sequence of rRNA bases are put
into the same group. Some of the new groups, such as Spirochetes, are essentially identical to early classication systems.
However, other groups contain a diverse assortment of bacteria that appear to be phenotypically distant from one another.
Gram-negative bacteria and Gram-positive bacteria are placed
in the same group as long as they share the same rRNA sequence of bases. The reading on the next page describes how
this methodology has led to the suggestion that there are three
domains of life, one of which includes the archaea, which were
formerly considered to be a type of bacteria.

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Archaea
Carl Woese at the University of Illinois has championed the hypothesis that sequencing of rRNA (ribosomal RNA) can be used
to classify organisms. He chose rRNA because of its involvement in protein synthesisits just possible that any changes
in rRNA sequence occur in a slow, steady manner as evolution
occurs.
On the basis of rRNA sequencing, Woese and his colleagues
concluded that there are three domains of life. A domain is a
higher classication category than the category kingdom. The
rst two domains, they maintain, are the Archaea and the Bacteria, both of which contain only unicellular prokaryotes. The
third domain, called Eukarya, contains all the eukaryotes (protists, fungi, plants, and animals).
Woeses hypothesis is remarkable because it suggests that
the archaea, long considered to be bacteria, should be considered as different from bacteria as bacteria are from eukaryotes.
The archaea are found in extreme environments thought to be
similar to those of the primitive earth (Figure 28A). The archaea
include the methanogens which live in anaerobic environments
such as swamps, marshes, and intestinal tracts of animals where
they produce methane (CH4) by a process that gives them energy for ATP formation. Methane produced by methanogenic
archaea is believed to contribute signicantly to global warming. The halophiles, another type of archaea, are salt lovers that
can be isolated from bodies of water like the Great Salt Lake in
Utah, the Dead Sea, solar salt ponds, and hypersalty soils. These
organisms pump chloride into their cells and synthesize ATP
only in the presence of light. The third major type of archaea are
the thermoacidophiles, which are both temperature and acid
loving. These archaea live in extremely hot, acidic environments
such as hot springs, geysers, submarine thermal vents, and
around volcanoes. They survive best at temperatures of about
80C, and some can grow even at 105C (remember that water
boils at 100C)! Many of the thermoacidophiles use sulfate as a
source of energy to make ATP and are found in natural environments with high sulfate concentrations.
Woese and his colleagues also went on to propose a new tree
of life. According to their rRNA-based tree, a universal common
ancestor gave rise to the Archaea and Bacteria and later, the Archaea gave rise to the Eukarya. This conclusion was based on
the nding that archaea have base sequences that are closer to
eukaryotic sequences than to bacterial sequences. But in science,
as you know, new information, often available because of new
technologies, may call into question former beliefs and hypotheses. Recently, new and more powerful sequencing instruments have allowed scientists to sequence the entire genomes of
bacteria and eukaryotic genomes, such as those of yeast. And
the whole genome data has called into question the tree of life
based on rRNA sequencing. The DNA genome data shows, to a
surprising degree, a mixture of DNA sequences from both archaea and bacteria in eukaryotes. For example, Russell Doolittle
who is with the Canadian Institute for Advanced Research in

Halifax, Nova Scotia, has found that 17 of 34 families of eukaryotic proteins look as if they came from bacteria and only eight
show a greater similarity to archaea, the supposed ancestor of
Eukarya. Even worse, some DNA data show a close relation between archaea and bacteria. Ron Swanson and Robert Feldman
of Diversa Corporation in San Diego found that the gene for an
enzyme involved in the synthesis of tryptophan , an amino acid,
is just about the same in Aquifex, domain Archaea, and Bacillus
subtilus, domain Bacteria. Robert Feldman has said, I think its
open whether the three domains [of life] will hold up.
Others have gone on to suggest the possibility of widespread
gene swapping among the rst organisms to evolve. In explanation, Doolittle says, You are what you eat, meaning that as
early unicellular organisms fed on one another they incorporated each others genes into their genomes. If gene swapping
occurred, Woese agrees that it will make it difcult to draw a
correct tree of life, but he still has faith that rRNA sequencing
data is sufcient to conclude that there are three domains of life:
Archaea, Bacteria, and Eukarya.

a.

b.

Figure 28A

Habitats of archaea.

a. Halophiles turn the water red in salt collection ponds in


California near San Francisco Bay. b. The scientic submarine
Alvin captures methanogen-containing material around a deepsea vent.

579

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a.

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28-8

Evolution and Diversity

;;;;
;;;;
;;;;
;;;;

b.

thylakoids

DNA

cell
wall

plasma
membrane

c.

storage granule

Figure 28.6 Diversity of the cyanobacteria.

a. In Gloeocapsa, single cells are grouped in a common gelatinous


sheath. b. Filaments of cells occur in Oscillatoria. c. One cell of
Oscillatoria as it appears through the electron microscope.

Cyanobacteria Cyanobacteria are gram-negative rods


with a number of unusual traits. They photosynthesize in
the same manner as plants and are believed to be responsible for rst introducing oxygen into the primitive atmosphere. Formerly, the cyanobacteria were called blue-green
algae and were classied with eukaryotic algae, but now we
know that they are prokaryotes. They can have other pigments that mask the color of chlorophyll, so that they appear, for example, not only blue-green but also red, yellow,
brown, or black.
Cyanobacterial cells are rather large and range in size
from 1 m to 50 m in length. They can be unicellular, colonial, or lamentous (Fig. 28.6). Cyanobacteria lack any visible means of locomotion, although some glide when in
contact with a solid surface and others oscillate (sway back
and forth). Some cyanobacteria have a special advantage

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because they possess heterocysts, which are thick-walled


cells without nuclei, where nitrogen xation occurs. The
ability to photosynthesize and also to x atmospheric nitrogen (N2) means they can obtain their nutritional requirements from the environment. They serve as food for
heterotrophs in aquatic ecosystems.
Cyanobacteria are common in fresh water, in soil, and
on moist surfaces, but they are also found in harsh habitats,
such as hot springs. They are symbiotic with a number of
organisms, such as liverworts, ferns, and even at times invertebrates like corals. In association with fungi, they form
lichens that can grow on rocks. Lichens help transform
rocks into soil. Other forms of life may then follow. The
rst moneran fossil dated to be 3.5 billion years old is believed to be a cyanobacterium, and there is evidence that
cyanobacteria were the rst colonizers of land during the
course of evolution.
Cyanobacteria are ecologically important in still
another way. If care is not taken in the disposal of nutrientrich industrial, agricultural, and human wastes, phosphates and nitrates drain into lakes and ponds, resulting in
a bloom of these organisms. The surface of the water becomes turbid, and light cannot penetrate to lower levels.
When a portion of the cyanobacteria die off, the decomposing bacteria use up the available oxygen, causing shes to
die from lack of oxygen.

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Cyanobacteria are photosynthesizers that


sometimes can also x atmospheric nitrogen. In
association with fungi, they form lichens, which are
important soil formers.

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Chapter 28

Reproduction of Bacteria

581

chromosome

The singular circular chromosome of bacteria consists only of double-stranded


DNA. Bacteria reproduce asexually by
means of binary ssion. First, the chromosome duplicates; then there are two chromosomes attached to the inside of the
plasma membrane. The chromosomes are
separated by an elongation of the cell,
which pushes the chromosomes apart.
Then the plasma membrane grows inward
and the cell wall forms, dividing the cell
into two daughter cells, each of which
now has its own chromosome (Fig. 28.7).
Sexual exchange of DNA occurs
among bacteria in three ways. Conjugation
takes place when the so-called male cell
passes DNA to the female cell by way of a
sex pilus. Transformation occurs when a
bacterium binds to and then takes up
DNA released into the medium by dead
bacteria. During transduction, bacteriophages carry portions of DNA from one
bacterium to another.
When faced with unfavorable environmental conditions, some bacteria form
endospores (Fig. 28.8). A portion of the cytoplasm and a copy of the chromosome dehydrate and are then encased by three
heavy, protective spore coats. The rest of
the bacterial cell deteriorates and the endospore is released. When environmental
conditions are again suitable for growth,
the endospore absorbs water and grows
out of the spore coats. In time, it becomes a
typical bacterial cell, capable of reproducing once again by binary ssion.

Microbiology

cell wall
plasma
membrane
cytoplasm

Figure 28.7 Binary ssion.


In electron micrographs, it is possible to observe a bacterium dividing to become two
bacteria. First DNA replicates, and as the plasma membrane lengthens, the two
chromosomes separate. Upon ssion, each bacterium has its own chromosome.

Figure 28.8 The endospore.


An endospore is resistant to extreme
environmental conditions. Sterilization, a process
that kills all living organismseven endospores
can be achieved by using an autoclave, a
container that maintains steam under pressure.
This bacterium, Bacillus subtilis, contains an
endospore.

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Killing Microorganisms
Viruses and bacteria are microbes that cause diseases in humans (Tables 28A and 28B). The development of drugs to kill
viruses has lagged far behind the development of those to kill
bacteria. Viruses lack most enzymes and, instead, utilize the
metabolic machinery of the host cell. Rarely has it been possible
to nd a drug that successfully interferes with viral reproduction without also interfering with host metabolism. One such
drug, however, called vidarabine, was approved in 1978 for
treatment of viral encephalitis, an infection of the nervous system. Acyclovir (ACV) seems to be helpful in treating genital herpes, and there are now various drugs (e.g., AZT) for the
treatment of AIDS. Since viral drugs are difcult to develop,
there is much concern about the possibility of other worldwide
epidemics as well as AIDS. The Ebola virus, which begins with
ulike symptoms and ends with vomiting and hemorrhaging, is
especially feared. Spread by direct contact with a victims blood
or other body uids, the disease is controllable only via strict
hygienic and sanitary controls.
An antibiotic is a drug that selectively kills bacteria. Most
antibiotics are produced naturally by soil microorganisms.
Penicillin is made by the fungus Penicillium, and streptomycin, tetracycline, and erythromycin are all produced by the
bacterium Streptomyces. Sulfa, a chemotherapeutic agent, is
produced in the laboratory. Antibiotics poison bacterial enzymes without harming host enzymes. Penicillin blocks the
synthesis of the bacterial cell wall; streptomycin, tetracycline,
and erythromycin block protein synthesis; and sulfa prevents
the production of a coenzyme. New antibiotics are being developed, but it will be some time before they are ready for
general use.

There are problems associated with antibiotic therapy. Some


patients are allergic to antibiotics, and the reaction can be fatal.
Antibiotics not only kill off disease-causing bacteria, they also
reduce the number of benecial bacteria in the intestinal tract
and other locations. These benecial bacteria hold in check the
growth of certain microbes that now begin to ourish. Diarrhea
can result, as can a vaginal yeast infection. The use of antibiotics
can also prevent natural immunity from occurring, leading to
the need for recurring antibiotic therapy. Most important, perhaps, is the growing resistance of certain strains of bacteria to
antibiotics. While penicillin used to be 100% effective against
hospital strains of Staphylococcus aureus, today it is far less effective. Tetracycline and penicillin, long used to cure gonorrhea,
now have a failure rate of more than 20% against certain strains
of gonococcus. Pulmonary tuberculosis is on the rise, particularly among AIDS patients, the homeless, and the rural poor,
and the new strains are resistant to the usual combined antibiotic therapy. A virulent streptococcal infection is now believed
to be the cause of the much publicized esh-eating condition
more properly called necrotizing fasciitis. About 30% of the people in the U. S. who develop fasciitis usually die.
To keep antibiotics effective, most physicians believe that they
should be administered only when absolutely necessary. Some
believe that if antibiotic use is not strictly limited, resistant strains
of bacteria will completely replace present strains and antibiotic
therapy will no longer be effective. They are much opposed to the
current practice of adding antibiotics to livestock feed in order to
make animals grow fatter. Bacteria that become resistant are easily transferred from animals to humans. Antibiotics have been a
boon to humans, but they should be used with care.

Table 28A Viral Diseases in Humans

Table 28B Bacterial Diseases in Humans

Category

Disease

Category

Disease

Sexually transmitted
diseases

AIDS (HIV), genital warts, genital herpes

Sexually transmitted
diseases

Syphilis, gonorrhea, chlamydia

Childhood diseases

Mumps, measles, chicken pox, German


measles

Respiratory diseases

Respiratory diseases

Common cold, inuenza, acute


respiratory infection

Strep throat, scarlet fever, tuberculosis,


pneumonia, Legionnaires disease,
whooping cough

Skin diseases

Erysipelas, boils, carbuncles,


impetigo, infections of surgical or
accidental wounds and burns, acne

Digestive tract diseases

Gastroenteritis, food poisoning,


dysentery, cholera

Nervous system diseases

Botulism, tetanus, spinal meningitis,


leprosy

Systemic diseases

Plague, typhoid fever, diphtheria

Other diseases

Gas gangrene, puerperal fever, toxic


shock syndrome, Lyme disease

Skin diseases

Warts, fever blisters, shingles

Digestive tract diseases

Gastroenteritis, diarrhea

Nervous system
diseases

Poliomyelitis, rabies, encephalitis

Other diseases

Cancer, hepatitis

582

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Chapter 28

28.3

583

Microbiology

Kingdom Protista

The protists are eukaryotes; their cells have a nucleus and all
of the various organelles. Unicellular organisms are predominant in kingdom Protista, and even the multicellular
forms lack the tissue differentiation that is seen in more
complex organisms. The protists are grouped according to
their mode of nutrition and other characteristics into the categories shown in the classication box. Three different types
of life cycles are typical of eukaryotes, and all three are seen
in kingdom Protista (Fig. 28.9). In the haplontic cycle, which
is typical of protists and fungi, the zygote is the only diploid
phase, and it undergoes meiosis to produce haploid spores.
In the alternation of generations cycle, which is typical of
plants, the sporophyte is a diploid individual that produces
spores by meiosis. In the diplontic cycle, which is typical of
animals, the diploid adult produces gametes by meiosis, and
the only haploid phase consists of the gametes.

Kingdom Protista
Eukaryotic; unicellular organisms and their immediate
multicellular descendants; sexual reproduction; agella and
cilia with 9 2 microtubules
Algae*
Phylum Chlorophyta: green algae
Phylum Phaeophyta: brown algae
Phylum Chrysophyta: diatoms and allies
Phylum Dinoagella: dinoagellates
Phylum Euglenophyta: euglenoids
Phylum Rhodophyta: red algae
Protozoans*
Phylum Sarcodina: amoebas and allies

Algae

Phylum Ciliophora: ciliates


Phylum Zoomastigophora: zooagellates

Algae are autotrophic by photosynthesis like plants. However, algae are aquatic, so they do not need to protect the zygote and embryo from drying out. Algae produce the food
that maintains communities of organisms in both the oceans
and bodies of fresh water. They are commonly named for the
type of pigment they contain; therefore, there are green,
golden brown, brown, and red algae. All algae contain chlorophyll, but they may also contain other pigments that mask the
color of the chlorophyll. Algae are grouped according to their
color and biochemical differences, such as the chemistry of
the cell wall and the way they store reserve food.

Phylum Sporozoa: sporozoa


Slime Molds*
Phylum Gymnomycota: slime molds
Water Molds*
Phylum Oomycota: water molds
*Categories which are not used in the classication of organisms, but added
here for clarity.

sporophyte

adult

(2n)

zygote
(2n)

(2n)
sporangium

diploid (2n)
meiosis

fertilization

zygote

zygote

haploid (n)
diploid (2n)
fertilization

meiosis
haploid (n)
diploid (2n)

spore

gametes

spore

meiosis

fertilization
haploid (n)

adult

gametophyte

(n)

(n)

a. Haplontic Cycle

b. Alternation of Generations

c. Diplontic Cycle

Zygote is 2n stage.
Meiosis produces spores.
Adult is always haploid.

Sporophyte is 2n generation.
Meiosis produces spores.
Gametophyte is haploid generation.

Adult is always 2n.


Meiosis produces gametes.

gametes

Figure 28.9 Common life cycles.

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28-12

Evolution and Diversity

Part 6

Green Algae
Green algae (phylum Chlorophyta, 7,000 species) live in the
ocean, but they are more likely found in fresh water and can
even be found on land, especially if moisture is available.
Some, however, have modications that allow them to live
on tree trunks, even in bright sun. Green algae are believed
to be closely related to the rst plants because both of these
groups (1) have a cell wall that contains cellulose, (2) possess
chlorophylls a and b, and (3) store reserve food as starch inside the chloroplast. (Other types of algae store reserve food
outside the chloroplast.) Green algae are not always green
because some have pigments that give them an orange, red,
or rust color.
Unicellular Green Algae Chlamydomonas is a unicellular
green alga usually less than 25 mm long that has two
whiplash agella (Fig. 28.10). A single, cup-shaped chloro-

zygote (2n)

zygospore (2n)
isogametes pairing
(n)

(n)

diploid (2n)

fertilization

meiosis
haploid (n)

plast contains a pyrenoid, where starch is synthesized. A


red-pigmented eyespot (stigma) is sensitive to light and
helps the organism detect light, which is necessary for photosynthesis.
When growth conditions are favorable, Chlamydomonas
reproduces asexually; the adult divides, forming zoospores
(agellated spores) that resemble the parent cell. A spore is a
reproductive cell that develops into a haploid individual
when environmental conditions permit. When growth conditions are unfavorable, Chlamydomonas reproduces sexually
according to the haplontic life cycle (see Fig. 28.9a). In most
species, the gametes are identical and are therefore called
isogametes. A heavy wall forms around the zygote, and it becomes a resistant zygospore able to survive until conditions
are favorable for germination. When a zygospore germinates, it produces four zoospores by meiosis.
Colonial Green Algae Volvox is a colony (loose association
of cells) in which thousands of agellated cells are arranged
in a single layer surrounding a watery interior. (Each cell of
a Volvox colony resembles a Chlamydomonas cell.) In Volvox,
the cells cooperate in that the agella beat in a coordinated
fashion. Cells that are specialized for reproduction divide
asexually to form a new daughter colony (Fig. 28.11). This
daughter colony resides for a time within the parental
colony, but then an enzyme that dissolves away a portion of
the parental colony allows it to escape. Sexual reproduction
among these algae involves heterogametes, that is, a denite
sperm and egg.

Sexual Reproduction

daughter
colony
gamete
formation

zoospores (n)

eyespot
nucleus

flagellum

chloroplast
starch
granule

zygote

pyrenoid

egg

Asexual Reproduction

zoospore (n)

zoospore
formation

sperm

Figure 28.10 Chlamydomonas, a agellated green alga.

Figure 28.11 Volvox, a colonial green alga.

During asexual reproduction, all structures are haploid. During sexual


reproduction, meiosis follows the zygote stage, which is the only
diploid part of the cycle. This is a haplontic life cycle (see Fig. 28.9a).

The adult Volvox colony often contains daughter colonies, which are
asexually produced by specialized cells. During sexual reproduction,
colonies produce a denite sperm and egg.

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Chapter 28

Filamentous Green Algae Filaments are end-to-end


chains of cells that form after cell division occurs in only one
plane. Spirogyra, a lamentous green alga, is found in green
masses on the surfaces of ponds and streams. It has chloroplasts that are ribbonlike and are arranged in a spiral within
the cell (Fig. 28.12). Conjugation, the temporary union of
two individuals during which there is an exchange of genetic material, occurs during sexual reproduction. The two
laments line up next to each other, and the cell contents of
one lament move into the cells of the other lament, forming diploid zygotes. These zygotes survive the winter, and
in the spring they undergo meiosis to produce new haploid
laments.
Multicellular Green Algae Multicellular Ulva is commonly called sea lettuce because of its leafy appearance (Fig.
28.13). The thallus is two cells thick and can be a meter long.
Ulva has an alternation of generations life cycle (see Fig.
28.9b) like that of plants, except that both generations look
exactly alike, the gametes look alike (isogametes), and the
spores are agellated.

585

Brown algae (phylum Phaeophyta, 1,500 species) range


from small forms with simple laments to large blade forms
between 50 meters and 100 meters in length (Fig. 28.14).
Large brown algae, often called seaweeds, are observed along
the rocky shoreline in the north temperate zone. These
plants are rmly anchored by holdfasts, and when the tide is
in, their broad attened blades are buoyed by air bladders.
When the tide is out, they do not dry up because their cell
walls contain a mucilaginous, water-retaining material.
Most brown algae have an alternation of generations life cycle, but some species of rockweed (Fucus) are unique in that
they have the diplontic life cycle (see Fig. 28.9c), in which
meiosis produces gametes and the adult is always diploid,
as in animals.
Brown algae provide food and habitat for marine organisms, and in several parts of the world they have been harsporophyte

Green algae can be unicellular, colonial,


lamentous, or multicellular. During sexual
reproduction, the zygote usually undergoes
meiosis and the adult is always haploid. Ulva has
an alternation of generations like plants do.

zygote

fertilization

Brown and Golden Brown Algae

Microbiology

diploid (2n)
haploid (n)

Brown and golden brown algae have chlorophylls a and c in


their chloroplasts and a type of carotenoid pigment (fucoxanthin) that gives them their color.

meiosis
plus (+)
gametophyte

+
spores
gametes

cell wall

nucleus

()
gametophyte

pyrenoid

Ulva Life Cycle

chloroplast

Figure 28.13 Ulva, a multicellular green alga.


Figure 28.12 Spirogyra, a lamentous green alga.
During conjugation the cell contents of one lament enter the cells of
another lament. Zygote formation follows.

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The sporophyte and gametophyte have the same appearance, and


the gametophyte produces isogametes. Ulva has an alternation of
generations life cycle, as do plants (see Fig. 28.9b).

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Part 6

28-14

Evolution and Diversity

Fucus

Sargassum

Laminaria

The cell wall of a diatom has an outer layer


of silica, a common ingredient of glass. The
valves are covered with a great variety of striations and markings, which form beautiful patterns when observed under the microscope.
Diatoms are among the most numerous of all
unicellular algae in the oceans. As such, they
serve as an important source of food for other
organisms. In addition, they produce a major
portion of earths oxygen supply. In ancient
times, diatoms were also present in astronomical numbers. Their remains, raised above sea
level by geological upheavals, are now mined
as diatomaceous earth for use as ltering agents,
soundproong materials, and scouring powders.

Dinoagellates
Many dinoagellates (phylum Dinoagella,
1,000 species) are bounded by protective celluLaminaria and Fucus are seaweeds known as kelps. They live along rocky coasts of the
lose plates (Fig. 28.15b). Most have two agella;
north temperate zone. Sargassum, the other brown alga shown, lives at sea where
one is free, but the other is located in a transoating masses form a home for many organisms.
verse groove. The beating of the agella causes
the organism to spin like a top. Occasionally,
vested for human food and for fertilizer. They are also a
when surface waters are warm and nutrients are high, there
source of algin, a pectinlike material that is added to ice
are so many of these unicellular organisms in the ocean that
cream, sherbet, cream cheese, and other products to give
they cause a condition called red tide. Toxins in red tides
them a stable, smooth consistency.
cause widespread sh kills and can paralyze humans who
Diatoms are a type of unicellular golden brown algae
eat shellsh that have fed on the dinoagellates.
(phylum Chrysophyta, 11,000 species). The structure of a diDinoagellates are an important source of food for small
atom is often compared to a box because the cell wall has
animals in the ocean. They also live as symbiotes within the
two halves, or valves, with the larger valve acting as a lid
bodies of some invertebrates. For example, because corals
for the smaller valve (Fig. 28.15a). When diatoms reproduce,
usually contain large numbers of dinoagellates, corals
each receives only one old valve. The new valve ts inside
grow much faster than they would otherwise.
the old one.

Figure 28.14 Diversication among the brown algae.

cellulose
plate

valve

flagella

a.

b.

c.

Figure 28.15 Diatoms and dinoagellates.


a. Diatoms may be variously colored, but their chloroplasts contain a unique golden brown pigment in addition to chlorophylls a and c.
The beautiful pattern results from markings on the silica-embedded wall. b. Dinoagellates have cellulose plates; these belong to Gonyaulax,
the dinoagellate that contains a red pigment and is responsible for occasional red tides.

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28-15

Chapter 28

Euglenoids
Euglenoids (phylum Euglenophyta, 1,000 species) are small
(10500 m) freshwater unicellular organisms that typify
the problem of classifying protists. One-third of all genera
have chloroplasts; the rest do not. Those that lack chloroplasts ingest or absorb their food. Euglenoids grown in the
absence of light have been known to lose their chloroplasts
and become heterotrophic. The chloroplasts are surrounded
by three rather than two membranes. The pyrenoid produces an unusual type of carbohydrate polymer (paramylon) not seen in green algae.
Euglenoids have two agella, one of which typically is
much longer than the other and projects out of an anterior
vase-shaped invagination (Fig. 28.16). It is called a tinsel agellum because it has hairs on it. Near the base of this agellum is an eyespot, which shades a photoreceptor for
detecting light. Because euglenoids are bounded by a exible pellicle composed of protein strips lying side by side, they
can assume different shapes as the underlying cytoplasm

long flagellum

photoreceptor

eyespot

short flagellum
contractile
vacuole

Microbiology

587

undulates and contracts. As in certain protozoans, there is a


contractile vacuole for ridding the body of excess water. Euglenoids reproduce by longitudinal cell division, and sexual
reproduction is not known to occur.
Euglenoids have both plant- and animal-like
characteristics. They have chloroplasts but lack a
cell wall and swim by agella.

Red Algae
Like the brown algae, red algae (phylum Rhodophyta, 4,000
species) are multicellular, but they live chiey in warmer seawater, growing in both shallow and deep waters. Red algae
are usually much smaller and more delicate than the brown
algae, although they can be up to a meter long. Some forms of
red algae are simple laments, but more often they are complexly branched, with the branches having a feathery, at, or
expanded ribbonlike appearance (Fig. 28.17). Coralline algae
are red algae that have cell walls impregnated with calcium
carbonate. In some instances, they contribute as much to the
growth of coral reefs as do coral animals.
Like brown algae, red algae are seaweeds of economic
importance. The mucilaginous material in the cell walls of
certain genera of red algae is a source of agar used commercially to make capsules for vitamins and drugs, as a material
for making dental impressions, and as a base for cosmetics.
In the laboratory, agar is a culture medium for bacteria.
When puried, it becomes the gel for electrophoresis, a procedure that separates proteins and nucleotides. Agar is also
used in food preparationas an antidrying agent for baked
goods and to make jellies and desserts set rapidly.
Many red algae have lamentous branches or are
multicellular.

nucleus
chloroplast
pyrenoid

nucleolus

pellicle

carbohydrate
granules

Figure 28.16 Euglena.


A very long agellum propels the body, which is enveloped by a
exible pellicle. A photoreceptor shaded by an eyespot allows
Euglena to nd light, after which photosynthesis can occur in the
numerous chloroplasts. Pyrenoids synthesize a reserve carbohydrate,
which is stored in the chloroplasts and also in the cytoplasm.

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Figure 28.17 Red alga.


Red algae are smaller and more delicate than brown algae.

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Part 6

28-16

Evolution and Diversity

Protozoans
Protozoans are typically heterotrophic, motile, unicellular
organisms of small size (21,000 m). They are not animals
because animals in the classication used by this text are
multicellular and undergo embryonic development.
Protozoans usually live in water, but they can also be
found in moist soil or inside other organisms. Some protozoans engulf whole food and are termed holozoic; others are
saprotrophic, and they absorb nutrient molecules across the
plasma membrane. Still others are parasitic and are responsible for several signicant human infections.

Most protozoans are unicellular, but they should not be


considered simple organisms. Each cell alone must carry out
all the functions performed by specialized tissues and organs in more complex organisms. They have organelles for
purposes we have not seen before. Their food is digested inside food vacuoles, and freshwater protozoans have contractile vacuoles for the elimination of water. Although
asexual reproduction involving binary ssion and mitosis is
the rule, many protozoans also reproduce sexually during
some part of their life cycle. The protozoans we will study
can be placed in four groups according to their type of locomotor organelle:
Name

pseudopod

contractile
vacuole

cytoplasm
food vacuole
a.

b.

Figure 28.18 Amoeboid protozoans.


a. Amoeba proteus is common in freshwater ponds. Bacteria and
other microorganisms are digested in food vacuoles, and contractile
vacuoles rid the body of excess water. b. Pseudopods of a live
foraminiferan project through holes in the calcium carbonate shell.
These shells were so numerous that they became a large part of the
White Cliffs of Dover when a geological upheaval occurred.

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Example

Amoeboids
Ciliates
Zooagellates
Sporozoa
nucleus

Locomotion
Pseudopods
Cilia
Flagella
No locomotion

Amoeba
Paramecium
Trypanosoma
Plasmodium

Amoeboids
The amoeboids (phylum Sarcodina, 40,000 species) are
protists that move and engulf their prey with
pseudopods. Amoeba proteus is a commonly studied freshwater member of this group (Fig. 28.18a). When amoeboids feed, they phagocytize; the pseudopods surround
and engulf the prey, which may be algae, bacteria, or other
protozoans. Digestion then occurs within a food vacuole.
Some white blood cells in humans are amoeboid, and they
phagocytize debris, parasites, and worn-out cells. Freshwater amoeboids, including Amoeba proteus, have contractile vacuoles where excess water from the cytoplasm
collects before the vacuole appears to contract, releasing
the water through a temporary opening in the plasma
membrane.
Entamoeba histolytica is a parasite that can infect the human intestine and cause amoebic dysentery. Complications
arise when this parasite invades the intestinal lining and reproduces there. If the parasites enter the body proper, liver
and brain impairment can be fatal.
The foraminifera, which are largely marine, have an external calcareous shell (made up of calcium carbonate)
with foramina, holes through which long, thin
pseudopods extend (Fig. 28.18b). The pseudopods branch
and join to form a net where the prey is digested.
Foraminifera live in the sediment of the ocean oor in incredible numbersthere may be as many as 50,000 shells
in a single gram of sediment. Deposits for millions of
years, followed by a geological upheaval, formed the
White Cliffs of Dover along the southern coast of England.
Also, the great Egyptian pyramids are built of
foraminiferan limestone. Today, oil geologists look for
foraminifera in sedimentary rock as an indicator of organic deposits, which are necessary for the formation of
oil.

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Chapter 28

Ciliates
The ciliates (phylum Ciliophora, 8,000 species) such as those
in the genus Paramecium are the most complex of the protozoans (Fig. 28.19). Hundreds of cilia, which beat in a coordinated rhythmic manner, project through tiny holes in a
semirigid outer covering, or pellicle. Numerous oval capsules
lying in the cytoplasm just beneath the pellicle contain trichocysts. Upon mechanical or chemical stimulation, trichocysts
discharge long, barbed threads, useful for defense and for capturing prey. When a paramecium feeds, food is swept down a
gullet, below which food vacuoles form. Following digestion,
the soluble nutrients are absorbed by the cytoplasm, and the
indigestible residue is eliminated at the anal pore.
During asexual reproduction, ciliates divide by transverse
binary ssion. Ciliates have two types of nuclei: a large
macronucleus and one or more small micronuclei. The macronucleus controls the normal metabolism of the cell; during sexual reproduction, two ciliates exchange a micronucleus.
The diversity of ciliates is quite remarkable. Barrelshaped didinia expand to consume paramecia much larger
than themselves. Suctoria have tentacles they use like straws

Microbiology

589

to suck their prey dry. Stentor looks like a blue vase decorated with stripes.

Zooflagellates
Protozoans that move by means of agella are called zooagellates (phylum Zoomastigophora, 1,500 species) to distinguish them from unicellular algae that also have agella.
Many zooagellates enter into symbiotic relationships. Trichonympha collaris lives in the gut of termites; it contains a bacterium that enzymatically converts the cellulose of wood to
soluble carbohydrates that are easily digested by the insect.
Giardia lamblia, whose cysts are transmitted through contaminated water, causes severe diarrhea. Trichomonas vaginalis, a
sexually transmitted organism, infects the vagina and urethra
of women and the prostate, seminal vesicles, and urethra of
men. A trypanosome, Trypanosoma brucei, transmitted by the
bite of the tsetse y, is the cause of African sleeping sickness
(Fig. 28.20). The white blood cells in an infected animal accumulate around the blood vessels leading to the brain and cut
off circulation. The lethargy characteristic of the disease is
caused by an inadequate supply of oxygen to the brain.

contractile vacuole
(partially full)

pellicle
cilia
food vacuoles
oral groove
macronucleus
micronucleus

flagellum

gullet
anal pore

contractile vacuole
(full)

a.

undulating
membrane

b.

Paramecium

Figure 28.19 Ciliated protozoans.

Back

Figure 28.20 Zooagellates.

Structure of Paramecium, adjacent to an electron micrograph.


Ciliates are the most complex of the protozoans. Note the oral
groove, the gullet, and the anal pore.

a. Photograph of Trypanosoma brucei, the cause of African sleeping


sickness, among red blood cells. b. The drawing shows its general
structure.

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28-18

Evolution and Diversity


zygote
salivary glands

female gamete

food canal

male gametes

salivary duct
2. When the mosquito
bites a human,
the sporozoites pass
from the mosquito
salivary glands into
the bloodstream and
then the liver of the host.

sporozoite

1. In the gut of female


Anopheles mosquito,
gametes fuse, and the
zygote undergoes many
divisions to produce
sporozoites, which
migrate to her salivary
gland.

3. Asexual spores
(merozoites) produced in
liver cells enter the
bloodstream and then
the red blood cells.
Liver
cell
4. When the red blood
cells rupture, spores
invade and reproduce
asexually inside new
red blood cells.

6. Some spores become male and


female gametocytes, which enter
the bloodstream. If taken up by a
mosquito, they become gametes.

Chills and
fever occur

Figure 28.21 Life cycle of Plasmodium

5. Spores and toxins pour


into the bloodstream
when the red blood cells
rupture.

vivax.
Asexual reproduction of Plasmodium occurs in
humans, while sexual reproduction takes place
within the Anopheles mosquito.

Sporozoa
Sporozoa (phylum Sporozoa, 3,600 species) are nonmotile
parasites of animals. Their name recognizes that these organisms form spores at some point in their life cycle.
Pneumocystis carinii causes the type of pneumonia seen
primarily in AIDS patients. The most widespread human
parasite is Plasmodium vivax, the cause of one type of
malaria. When a human is bitten by an infected female
Anopheles mosquito, the parasite eventually invades the red
blood cells. The chills and fever of malaria appear when the
infected cells burst and release toxic substances into the
blood (Fig. 28.21). Malaria is still a major killer of humans,
despite extensive efforts to control it. A resurgence of the
disease was caused primarily by the development of
insecticide-resistant strains of mosquitoes and by parasites
resistant to current antimalarial drugs.
The protozoans are the animal-like protiststhey
ingest their food and are motile. Protozoans are
classied according to the type of locomotor
organelle employed.

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Slime Molds and Water Molds


Slime molds (phylum Gymnomycota, 560 species) might
look like molds, but their vegetative state is amoeboid,
whereas fungi are lamentous. Fungi are saprotrophic,
whereas slime molds are heterotrophic by ingestion. When
conditions are unfavorable to growth, however, slime molds
produce and release spores that are resistant to environmental extremes. Fungi also produce such spores.
Usually plasmodial slime molds exist as a plasmodium, a
diploid multinucleated cytoplasmic mass enveloped by a
slime sheath that creeps along, phagocytizing decaying
plant material in a forest or agricultural eld (Fig. 28.22).
Under unfavorable conditions, the plasmodium forms sporangia, structures that produce spores which are dispersed
by the wind. The spores germinate to produce gametes that
join to form a zygote. This zygote begins the cycle again. Cellular slime molds, as you might expect, exist as individual
amoeboid cells. Each lives by phagocytizing bacteria and
yeast. As the food supply runs out, the cells release a chemical that causes them to aggregate into a pseudoplasmodium
that produces spores within sporangia.

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Chapter 28

Water molds (phylum Oomycota, 580 species) live in the


water, where they parasitize sh, forming furry growths on
their gills. Others live on land and parasitize insects and
plants; a water mold was responsible for the 1840s potato
famine in Ireland. Most water molds, like fungi, are saprotrophic and have a lamentous body, but they have the
diplontic life cycle (see Fig. 28.9c), whereas fungi have the
haplontic cycle (see Fig. 28.9a).

28.4

Microbiology

591

Kingdom Fungi

Fungi are multicellular eukaryotes that are heterotrophic by


absorption. They send out digestive enzymes into the immediate environment, and then, when organic matter is broken down, they absorb nutrient molecules. Like bacteria,
most fungi are saprotrophic decomposers that break down the
waste products and dead remains of plants and animals.
Some fungi parasitize both plants and animals; in humans,
they cause ringworm, athletes foot, and yeast infections.
Although yeast are unicellular fungi, the body of a fungus is usually a multicellular structure known as a
mycelium. A mycelium is a network of laments called hyphae (sing., hypha):

septum

Hypha

cell wall

nuclei

Kingdom Fungi
Multicellular eukaryote; heterotrophic by absorption; lack
agella; nonmotile spores form during both asexual and
sexual reproduction
Division Zygomycota: zygospore fungi
Soil and dung molds, black bread molds (Rhizopus).
Division Ascomycota: sac fungi
Many small wood-decaying fungi, yeasts (Saccaromyces),
molds (Neurospora), morels, cup fungi, trufes; plant
parasites: powdery mildews, ergots.
Division Basidiomycota: club fungi
Mushrooms, stinkhorns, puffballs, bracket and shelf fungi,
coral fungi; plant parasites: rusts, smuts.
Division Deuteromycota: imperfect fungi (i.e., means of
sexual reproduction not known)
Athletes foot, ringworm, candidiasis.

go

po

zy

res

meiosis

te

g a m ete

Figure 28.22 Plasmodium and life cycle of a yellow slime


mold, Hemitrichia stipitata.

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Fungal cells are quite different from plant cells not only by
lacking chloroplasts but also by having a cell wall that contains chitin and not cellulose. Chitin is a polymer of glucose,
but each glucose molecule has an amino group attached to
it. (Chitin is also found in the external skeleton of insects and
all arthropods.) How can a nonmotile terrestrial organism
ensure that the species will be dispersed to new locations?
Fungi produce nonagellate spores during both sexual and
asexual reproduction, which are dispersed by the wind.

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Evolution and Diversity

thick-walled
zygospore

zygote

diploid (2n)

nuclear
fusion

meiosis
sporangium

haploid (n)

Sexual
Reproduction
spores (n)

gametangia

cytoplasmic
fusion

+ mating type
zygospore
germination

germination
of spores

mating type
spores (n)

Asexual
Reproduction

sporangium
sporangiophore
mycelium

5 m
stolon
rhizoid

mating type

+ mating type

Figure 28.23 Black bread mold, Rhizopus stolonifer.


Asexual reproduction is the norm. During sexual reproduction, two compatible mating types make contact: rst gametangia fuse, and then nuclei
fuse. The zygospore is a resting stage that can survive unfavorable growing conditions. Meiosis occurs when the zygote germinates and spores
are released from a sporangium.

Zygospore Fungi
The zygospore fungi (phylum Zygomycota, 600 species) live
off plant and animal remains in the soil and also bakery
goods in our kitchens. Some, however, are parasites of small
soil protists or worms, and even insects such as the housey.
In Rhizopus stolonifer, black bread mold, stolons are horizontal hyphae that exist on the surface of the bread; rhizoids
grow into the bread, and sporangiophores are stalks that bear
sporangia (Fig. 28.23). A sporangium is a capsule that produces spores, more properly called sporangiospores. During
asexual reproduction all structures involved are haploid;
during sexual reproduction there is a diploid zygospore for
which the phylum is named. Hyphae of opposite mating

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types, termed plus () and minus (), grow toward each


other until they touch. Gametangia form and merge, producing a large cell in which nuclei of the two mating types pair
and then fuse. A thick wall develops around the cell, which
is now called a zygospore. Upon germination, sporangiophores develop, and many spores are produced by meiosis.
By now, you will have no trouble in identifying this cycle as
a haplontic life cycle (see Fig. 28.9a).

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Zygospore fungi produce spores within sporangia.


During sexual reproduction, a zygospore forms
prior to meiosis and production of spores.

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Chapter 28

Microbiology

593

b.

ascospores
a.

zygote
nuclear (2n)
fusion

mature
ascus
meiosis

Figure 28.24 Sac fungi.


a. Fruiting body of a cup
fungus. Nuclear fusion and
zygote formation lead to a
mature ascus, where
ascospores are produced. b.
Although yeast on occasion
produce ascospores, they
usually reproduce by budding,
as shown here.

dikaryotic hyphae
male organ
female organ
+ mating type (n)
spore

mating type (n)


spore

Sac Fungi

Yeasts

Sac fungi (division Ascomycota, 30,000 species) include red


bread molds (e.g., Neurospora) and cup fungi (Fig. 28.24a).
Also, morels and trufes are sac fungi highly prized as
gourmet delicacies. A large number of sac fungi are parasitic
on plants; powdery mildews grow on leaves, as do leaf curl
fungi; chestnut blight and Dutch elm disease destroy the
trees named. Ergot is a parasitic sac fungus that infects rye
and (less commonly) other grains.
The division name for sac fungi, Ascomycota, refers to
the ascus, a ngerlike sac that develops after hyphae from
two mating strains merge, producing dikaryotic (each cell
has two nuclei) hyphae. In an ascus, a zygote forms and undergoes meiosis to produce eight haploid nuclei that become
eight ascospores.
ascospore

ascus

The asci are usually surrounded and protected by sterile


hyphae within a fruiting body. Asexual reproduction,
which is the norm among ascomycetes, involves the production of spores called conidiospores (see Fig. 28.26).

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Yeasts are unicellular sac fungi that reproduce asexually either by mitosis or by budding (Fig. 28.24b). When yeasts ferment, they produce ethanol and carbon dioxide. In the wild,
yeasts grow on fruits, and historically the yeasts already present on grapes were used to produce wine. Today selected
yeasts are added to relatively sterile grape juice in order to
make wine. Also, yeasts are added to prepared grains to
make beer. Both the ethanol and the carbon dioxide are retained for beers and sparkling wines; carbon dixoide is released for still wines. In baking, the carbon dioxide given off
by yeast is the leavening agent that causes bread to rise.
Yeasts are serviceable to humans in another way. They
have become the material of choice in genetic engineering
experiments requiring a eukaryote. Escherichia coli, the usual
experimental material, is a prokaryote and does not function
during protein synthesis as a eukaryote would.
When sac fungi reproduce sexually, they produce
ascospores within asci, usually within a fruiting
body.

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Evolution and Diversity

Club Fungi
Club fungi (phylum Basidiomycota, 16,000 species) include
shelf or bracket fungi on dead trees and mushrooms in
lawns and forests. Less well known are puffballs, birds nest
fungi, and stinkhorns. These structures are all fruiting bodies that contain basidia, club-shaped structures that give
this phylum its name.
Club fungi usually reproduce sexually (Fig. 28.25). Hyphae from two different mating types meet, and cytoplasmic
fusion occurs. The resulting dikaryotic mycelium periodically produces fruiting bodies, which are composed of
tightly packed hyphae. The fruiting body of a mushroom

has a stalk and a cap. The cap of a gilled mushroom contains


radiating lamellae lined by basidia where nuclear fusion,
meiosis, and spore production occur. A basidium has four
projections into which cytoplasm and a haploid nucleus enter. The spores are windblown and germinate to give a haploid mycelium.

Club fungi usually reproduce sexually. The


dikaryotic stage is prolonged and periodically
produces fruiting bodies where spores are
produced in basidia.

basidium

zygote
diploid (2n)

nuclear fusion

meiosis

nuclei

dikaryotic
(n + n)

basidium

basidiospores

monokaryotic
(n)

spores are released

portion of gill

cap

monokaryotic
mycelia

gill

germination of spores

dikaryotic
mycelium

stalk

a.

cytoplasmic
fusion

fruiting body
(basidiocarp)

b.

Figure 28.25 Club fungi.


a. Life cycle of a mushroom in which sexual reproduction is the norm. After hyphae from two compatible mating types fuse, the dikaryotic
mycelium is long-lasting. Nuclear fusion results in zygotes within basidia on the gills of the fruiting body shown. Meiosis occurs and
basidiospores are released. b. Fairy ring. Mushrooms develop in a ring on the outer living fringes of a dikaryotic mycelium. The center has used
up its nutrients and is no longer living.

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Chapter 28

conidiospore

Figure 28.26 Conidiospores.


Sac fungi and imperfect fungi reproduce asexually by producing
spores called conidiospores at the ends of certain hyphae. The
organism shown here is Penicillium, an imperfect fungus.

Microbiology

595

Lichens are a symbiotic relationship between a fungus


and a cyanobacterium or a green alga. The body of a lichen
has three layers: the fungus forms a thin, tough upper layer
and a loosely packed lower layer that shield the photosynthetic cells in the middle layer (Fig. 28.27a). In the past,
lichens were assumed to be a relationship of mutual benet:
the fungus received nutrients from the algal cells, and the algal cells were protected from desiccation by the fungus. Actually, lichens may involve a controlled form of parasitism of
the algal cells by the fungus and algae may not benet at all
from these associations. This is supported by experiments in
which the fungal and algal components are removed and
grown separately. It is difcult to cultivate the fungus,
which does not usually grow alone.
Three types of lichens are recognized. Compact crustose
lichens (Fig. 28.27b) are often seen on bare rocks or on tree
bark; foliose lichens are leaike; and fruticose lichens are
shrublike. Lichens can live in areas of extreme environmental
conditions and are important soil formers. In Arctic ecosystems, a lichen called reindeer moss is a common photosynthetic organism and an important food source for animals.

Imperfect Fungi
The imperfect fungi (phylum Deuteromycota, 25,000
species) always reproduce asexually by forming conidiospores (Fig. 28.26). These fungi are imperfect in the sense
that no sexual stage has yet been observed and may not exist. Without knowing the sexual stage, it is often difcult to
classify a fungus as belonging to one of the other phyla.
Several imperfect fungi are serviceable to humans.
Some species of the mold Penicillium are sources of the antibiotic penicillin, while other species give the characteristic flavor and aroma to cheeses such as Roquefort and
Camembert. The bluish streaks in blue cheese are patches
of conidiospores. The drug cyclosporine, which is administered to suppress the immune system following an organ
transplant operation, is derived from an imperfect fungus
found in soil.
Unfortunately, some imperfect fungi cause disease in
humans. Certain dust-borne spores can cause infections of
the respiratory tract, while athletes foot and ringworm are
spread by direct contact. Candida albicans is a yeastlike organism that causes infections of the vagina, especially in
women on the birth-control pill. This organism also causes
thrush, an inammation of the mouth and throat.

reproductive unit
fungal hyphae

algal cells

algal layer

fungal hyphae

a.

The imperfect fungi cannot be classied into one


of the other phyla because their mode of sexual
reproduction is unknown.
b.

Fungal Relationships

Figure 28.27 Lichen morphology.

We have already mentioned several instances in which fungi


are parasites of plants and animals. Two other associations
are of interest.

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a. A section of a lichen shows the placement of the algal cells and


the fungal hyphae, which encircle and penetrate the algal cells.
b. Mixture of compact crustose lichens.

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Evolution and Diversity

Mycorrhizae (fungus roots) are symbiotic relationships


of mutual benet between soil fungi and the roots of most
plants. The fungus provides the plants with inorganic nutrients, and the plant provides the fungus with organic nutrients. Plants whose roots are infected with mycorrhizae grow

arriers of disease are persons who do


not appear to be ill but who can
nonetheless pass on an infectious disease.
Carriers of sexually transmitted diseases
can pass on an infection to their partners,
and even sometimes to people who have
never had sexual relations with them. Hepatitis C and HIV are transmitted by blood
to blood contact, as when drug abusers
share needles.
The only way society can protect itself
is to identify carriers and remove them
from areas or activities where transmission of the pathogen is most likely. Sometimes its difcult to identify activities that
might pass on a pathogen like HIV. A few
people believe that they have acquired
HIV from their dentists, and while this is
generally believed to be unlikely, medical
personnel are still required to identify

themselves when they are carriers of HIV.


Magic Johnson is a famous basketball star
who is HIV positive. When he made it
known that he was a carrier, some of his
teammates refused to play basketball with
him and he had no choice but to retire.
Later, Johnson did play again for a while.
There are other sports in which transmission of HIV is more likely. The Centers for
Disease Control did a statistical study to
try to gure the odds of acquiring HIV
from another football player. They gured
that the odds were 1 in 85 million.
The odds might be higher for boxing,
a bloody sport. When two brothers, one of
whom had AIDS, got into a vicious ght,
the infected brother repeatedly bashed his
head against his brothers. Both men bled
profusely and soon after, the previously
uninfected brother tested positive for the

Summarizing the Concepts


28.1 Viruses
Viruses are noncellular obligate parasites that have a protein coat
called a capsid and a nucleic acid core. Viral DNA must enter a host cell
before reproduction is possible. In the lytic cycle, a bacteriophage immediately reproduces, and in the lysogenic cycle, viral DNA integrates
into the host genome and may eventually reproduce.

28.2 Kingdom Monera


The kingdom Monera includes prokaryotic unicellular organisms,
namely the bacteria. Most bacteria are saprotrophs (heterotrophic by
absorption), and along with fungi, fulfill the role of decomposers
in ecosystems. The cyanobacteria are photosynthetic in the same
manner as plants. Reproduction of bacteria is by binary ssion, but
sexual exchange occasionally takes place. Some bacteria form
endospores, which can survive the harshest of treatment except
sterilization.

28.3 Kingdom Protista


Kingdom Protista includes eukaryotic unicellular organisms and some
multicellular forms. Algae are aquatic autotrophs by photosynthesis;
protozoans are aquatic heterotrophs by ingestion. Slime molds, which
are terrestrial, and water molds, which are aquatic, have some characteristics of fungi.

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more successfully in poor soilsparticularly soils decient


in phosphatesthan do plants without mycorrhizae. The
fungal partner may enter the cortex of roots but does not enter plant cells.

virus. The possibility of transmission of


HIV in the boxing ring has caused several
states to require boxers to undergo routine
HIV testing. If they are HIV positive, they
cant ght.

Questions
1.

2.

3.

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Should all people who are HIV positive


identify themselves? Why or why not? By
what method would they identify
themselves at school, at work, and other
places?
What type of contact would you have
with an HIV positive individual? Would
you play on the same team? Would you
have sexual relations with an HIV positive
individual if a condom was used?
Discuss.
Is it ethical to make an HIV positive
individual feel stigmatized? Discuss.

Algae are classied according to their pigments (colors). Green algae are diverse: some are unicellular or colonial agellates, some are
lamentous, and some are multicellular sheets. Diatoms and dinoagellates are unicellular producers in oceans; brown and red algae are
seaweeds; euglenoids are unicellular with both plant and animal characteristics. Every type of life cycle is seen among the algae.
Protozoans are classied according to the type of locomotor organelle. The amoebas phagocytize, the ciliates are very complex, and
sporozoa are all animal parasites. Malaria is a signicant disease
caused by a sporozoan.
Slime molds have an amoeboid stage and then form fruiting bodies, which produce spores that are dispersed by the wind. Water molds
have threadlike bodies.

28.4 Kingdom Fungi


Kingdom Fungi includes eukaryotic multicellular organisms that are
saprotrophs. Fungi are composed of hyphae, which form a mycelium.
Along with heterotrophic bacteria, they are decomposers. The fungi
produce windblown spores during both sexual and asexual reproduction. The major groups of fungi are distinguished by type of sexual
spore and fruiting body. Zygospore fungi produce spores in sporangia;
sac fungi produce ascospores in asci; and club fungi form basidiospores in basidia.
Fungi form two symbiotic associations of interest. Lichens contain
both a fungus and an alga; mycorrhizae is a symbiotic relationship of
mutual benet between soil fungi and roots of plants.

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Chapter 28

Studying the Concepts


1. Describe the structure of viruses and the manner of reproduction of bacteriophages, including the lytic and lysogenic
cycles. 57475
2. Describe the life cycle of a retrovirus and explain why retroviruses are of interest today. 576
3. In general, describe the structure of prokaryotic cells. 577
4. Describe the metabolic diversity of bacteria and how they are
classied. 57780
5. Describe the manner in which bacteria reproduce, exchange
genetic material, and form endospores. 581
6. What are the three groups of protists and how is each group
distinguishable? 583
7. Describe the different types of green algae and how their
means of sexual reproduction differ. Show that the life cycle
of Chlamydomonas is an example of the haplontic cycle and
that the life cycle of Ulva is an example of the alternation of
generations life cycle. 58485
8. In what ways are brown and red algae similar? How are they
different? 58587
9. In what ways are diatoms and dinoagellates similar? How
are they different? 586
10. What are the animal-like characteristics of euglenoids? The
plantlike characteristics? 587
11. Why are the slime molds and the water molds sometimes
called the funguslike protists? 59091
12. Describe the anatomical features of fungi, and tell how fungi
are classied. 591
13. Describe the life cycles of black bread mold and a
mushroom. 592, 594
14. Describe the structure of lichens and the importance of both
lichens and mycorrhizae. 59596

Testing Yourself
Choose the best answer for each question.
1. Which of these are found in all viruses?
a. envelope, nucleic acid, capsid
b. hyphae and cilia
c. DNA, RNA, and proteins
d. proteins and a nucleic acid
e. proteins, nucleic acids, carbohydrates, and lipids
2. Which step in the lytic cycle follows attachment of the virus
and release of DNA into the cell?
a. production of lysozyme
b. biosynthesis of viral components
c. assemblage
d. integration of viral DNA into host DNA
e. host DNA replication
3. RNA retroviruses have a special enzyme that
a. disintegrates host DNA.
d. translates host DNA.
b. polymerizes host DNA.
e. repairs viral DNA.
c. transcribes viral RNA to cDNA.
4. Facultative anaerobes
a. require a constant supply of oxygen.
b. are killed in an oxygenated environment.
c. do not always need oxygen.
d. are photosynthetic.
e. are chemosynthetic.

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Microbiology

597

5. Cyanobacteria, unlike other types of bacteria that photosynthesize, do


a. not give off oxygen.
d. not have a cell wall.
b. give off oxygen.
e. form plasmodia.
c. not have chlorophyll.
6. Chemosynthetic bacteria
a. are autotrophic.
b. use the rays of the sun to acquire energy.
c. oxidize inorganic compounds to acquire energy.
d. Both a and c are correct.
e. Both a and b are correct.
7. Which is mismatched?
a. red algaemulticellular, delicate, seaweed
b. diatomssilica shell, boxlike, golden brown
c. euglenoidsagella, pellicle, eyespot
d. Fucusadult is diploid, seaweed, chlorophylls a and c
e. Parameciumcilia, calcium carbonate shell, gullet
8. Which is a false statement?
a. Slime molds and water molds are protists.
b. There are agellated algae and agellated protozoans.
c. Among protozoans, both agellates and sporozoans are
symbiotic.
d. Among protists, only green algae have a sexual life
cycle.
e. Ciliates exchange genetic material during
conjugation.
9. Which is mismatched?
a. water moldpotato famine
b. trypanosomeAfrican sleeping sickness
c. Plasmodium vivaxmalaria
d. amoeboidsevere diarrhea
e. AIDSGiardia lamblia
10. Which is found in slime molds but not fungi?
a. nonmotile spores
d. photosynthesis
b. amoeboid adult
e. All of these are correct.
c. zygote formation
11. The taxonomy of fungi is based on
a. sexual reproductive structures.
b. shape of the sporocarp.
c. mode of nutrition.
d. type of cell wall.
e. All of these are correct.
12. In the life cycle of black bread mold, the zygospore
a. undergoes meiosis and produces zoospores.
b. produces spores as a part of asexual reproduction.
c. is a thick-walled dormant stage.
d. is equivalent to asci and basidia.
e. Both a and c are correct.
13. When sac fungi and club fungi reproduce sexually, they
produce
a. a fruiting body.
b. spores.
c. conidiospores.
d. hyphae.
e. Both a and b are correct.
14. Lichens
a. are comprised of bacteria and fungi.
b. cannot reproduce.
c. need a nitrogen source to live.
d. are parasitic on trees.
e. are able to live in extreme environments.

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Evolution and Diversity

Part 6

15. Label this diagram of the Chlamydomonas life cycle.

Understanding the Terms

c.

alga (pl., algae) 583


alternation of
generations 585
archaea 578
ascus 593
bacteriophage 575
bacterium 577
basidium 594
binary ssion 581
ciliate 589
conidiospore 593
conjugation 585
cyanobacterium 580
decomposer 578
diatom 586
dinoagellate 586
diplontic life cycle 585
lament 585
fruiting body 593
fungus (pl., fungi) 591
haplontic life cycle 584
hypha 591

d.

b.
(n)

(n)

diploid (2n)
meiosis

fertilization

haploid (n)

a.

m.
f.
l.
g.

k.

h.
j.

lichen 595
lysogenic cycle 575
lytic cycle 575
mad cow disease 577
mycelium 591
mycorrhiza 596
nucleoid 577
phagocytize 588
plasmodium 590
prion 577
protozoan 588
pseudopod 588
retrovirus 576
saprotroph 578
sporangium 592
spore 584
symbiotic 578
trypanosome 589
viroid 577
virus 574
zooagellate 589
zygospore 592

i.

e.

zoospore (n)

zoospore
formation

n. Give two reasons why each portion of the cycle is either


asexual or sexual.

Match the terms to these denitions:


a.
Portion of the bacterial cell that contains its
genetic material.
b.
Relationship that occurs when two different
species live together in a unique way; it may be benecial,
neutral, or detrimental to one and/or the other species.
c.
Splitting of a parent cell into two daughter cells;
serves as an asexual form of reproduction in bacteria.
d.
Organism that secretes digestive enzymes and
absorbs the resulting nutrients back across the plasma membrane.
e.
Symbiotic relationship between certain fungi and
algae, in which the fungi possibly provide inorganic food or
water and the algae provide organic food.

Thinking Scientically
1. In reference to the life cycles (Figure 28.9):
a. The timing of what one element determines whether or not
a diploid adult results?
b. Hypothesize how the haplontic life cycle may have given
rise to the alternation of generations life cycle.
c. Hypothesize how the alternation of generations life cycle
may have given rise to the diplontic life cycle.

Using Technology
Your study of microbiology is supported by these available
technologies.

2. Members of the kingdom Fungi are adapted to living on land.


a. What characteristic of fungi would make you think that
other organisms (such as plants and animals) must have
been present on land before fungi?
b. Both Chlamydomonas and fungi follow the haplontic life
cycle. What elements in the cycle show that the former is
adapted to living in the water and the latter is adapted to
living on land?
c. The mycelia of sac fungi and club fungi are found in the
ground, and the fruiting bodies usually appear after a rain.
What does this tell you about the adaptation of fungi to
living on land?

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Essential Study Partner CD-ROM


Evolution & Diversity Viruses
Bacteria
Protists
Fungi
Visit the Mader web site for related ESP activities.

Exploring the Internet


The Mader Home Page provides resources and tools as
you study this chapter.

http://www.mhhe.com/biosci/genbio/mader

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