MICROTAXONOMY 04
form endospores.
Domain Bacteria: the NON-Proteobacteria
I. The Gram-Positive Bacteria
 Gram-positive bacteria were historically grouped
based on their general shape (e.g., rods, cocci, or
irregular) and their ability to form endospores.
 Analysis of the phylogenetic relationships within the
gram-positive bacteria by comparison of 16S rRNA
sequences shows that they are divided into a low G
+ C group and high G + C, or actinobacterial, group.
A. FIRMICUTES: The Low G+C Gram-positives
 Bergey’s Manual groups the gram-positive bacteria
phylogenetically into two major groups: the low G +
C gram-positive bacteria and the high G + C gram-
positive bacteria.
- This classification is based primarily on nucleic acid
sequences rather than phenotypic similarity.
 The low G + C gram positives contain (1) clostridia
and relatives, (2) the mycoplasmas, and (3) the
bacilli and lactobacilli. Endospore formers, cocci, and
rods are found among the clostridia and bacilli
groups.
- Thus, common possession of a complex structure
such as an endospore does not necessarily indicate
close relatedness between the genera.
 Peptidoglycan structure varies among different
groups in ways that are often useful in their
identification.
 Most gram-positive bacteria are harmless, free-living
saprophytes, but most major groups include
pathogens of humans, other animals, and plants.
- Some gram-positive bacteria are very important in
the food and dairy industries.
Genus Clostridium
 By far the largest genus is Clostridium (klôs-triʹdē-
um) under Order Clostridiales. It includes obligately
anaerobic, fermentative, gram-positive bacteria that
C. acetobutylicumis used to manufacture butanol in
- The genus contains well over 100 species in several
distinct phylogenetic clusters. The genus Clostridium
may be subdivided into several genera in the future.
 Members of the genus Clostridium have great
practical impact. Because they are anaerobic and
form heat-resistant endospores, they are
responsible for many cases of food spoilage, even in
canned foods.
 Clostridia often can ferment amino acids to produce
ATP by oxidizing one amino acid and using another
as an electron acceptor in a process called the
Stickland reaction.
- This reaction generates ammonia, hydrogen sulfide,
fatty acids, and amines during the anaerobic
decomposition of proteins. These products are
responsible for many unpleasant odors arising
during putrefaction.
 C. difficile (difʹfi-sē-il) is an inhabitant of the
intestinal tract that may cause a serious diarrhea.
This occurs only when antibiotic therapy alters the
normal intestinal microbiota, allowing overgrowth
by toxin producing C. difficile.
 Several clostridia produce toxins and are major
disease agents. Diseases associated with clostridia
include tetanus, caused by C. tetani (teʹtan-e);
botulism, caused by C. botulinum (bo-tū-līʹnum); and
gas gangrene , caused by C. perfringens (per-
frinʹjens) and other clostridia. C. perfringens is also
the cause of a common form of foodborne diarrhea,
gas gangrene and food poisoning.
- C. perfringens genome sequence analysis reveals
that the microbe possesses the genes for
fermentation with gas production but lacks genes
encoding enzymes for the TCA cycle or a respiratory
chain.
- C. perfringens has an extraordinary doubling time of
only 8 to 10 minutes when in the human host.
Clostridia also are industrially valuable; for example,
some countries.
SEE FIGURE 1
Genus Epulopicium
 Biologists have long considered bacteria to be small
by necessity because they lack the nutrient
transport systems used by higher, eukaryotic
organisms and because they depend on simple
diffusion to obtain nutrients. These characteristics
would seem to critically limit size. So, when a cigar-
shaped organism living symbiotically in the gut of
the Red Sea surgeonfish was first observed in 1985,
it was considered to be a protozoan.
 Its size suggested this: the organism was as large as
80 μm ×600 μm— over half a millimeter in length—
large enough to be seen with the unaided eye.
Compared to the familiar bacterium E. coli, which is
about 1 μm ×2 μm, this organism would be about a
million times larger in volume.
 Further investigation of the new organism showed
that certain external structures thought to resemble
the cilia of protozoa were actually similar to
bacterial flagella, and it did not have a membrane-
enclosed nucleus.
 Ribosomal RNA analysis conclusively placed
Epulopiscium (epʹū-lō-pis-ē-um) with the
prokaryotes. (The name means “guest at the
banquet of a fish.” It is literally bathed in semi
digested food.)
 It most closely resembles gram-positive bacteria of
the genus Clostridium.
- Strangely, the species Epulopiscium fishelsoni
(fishel-sōʹnē) does not reproduce by binary fission.
Daughter cells formed within the cell are released
through a slit opening in the parent cell. This may be
related to the evolutionary development of
sporulation.
Genus Bacillus
 Bacteria of the genus Bacillus are typically rods that
produce endospores. They are common in soil, and
only a few are pathogenic to humans. Several
species produce antibiotics.
 Bacillus anthracis (bä-silʹlusan-thrāʹsis) causes
anthrax, a disease of cattle, sheep, and horses that
can be transmitted to humans.
- It is often mentioned as a possible agent of
biological warfare. The anthrax bacillus is a
nonmotile facultative anaerobe, often forming
chains in culture. The centrally located endospore
does not distend the walls.
 Bacillus thuringiensis (thur-in-jē-enʹsis) is probably
the best-known microbial insect pathogen. It
produces intracellular crystals when it sporulates.
Commercial preparations containing endospores
and crystalline toxin (Bt) of this bacterium are sold
in gardening supply shops to be sprayed on plants.
 Bacillus cereus (seʹrē-us) is a common bacterium in
the environment and occasionally is identified as a
cause of food poisoning, especially in starchy foods
such as rice.
 The three species of the genus Bacillus that we have
just described are dramatically different in
important ways, especially their disease-causing
properties. However, they are so closely related that
taxonomists consider them to be variants of a single
species, differing almost entirely in genes carried on
plasmids, which are easily transferred from one
bacterium to another.
Genus Staphylococcus
 Staphylococci typically occur in grape-like clusters.
The most important staphylococcal species is
Staphylococcus aureus (staf-i-lō-kokʹkusôʹrē-us),
which is named for its yellow pigmented colonies
(aureus = golden). Members of this species are
facultative anaerobes.
 Some characteristics of the staphylococci account
for their pathogenicity, which takes many forms.
 They grow comparatively well under conditions of
high osmotic pressure and low moisture, which
partially explains why they can grow and survive in
nasal secretions (many of us carry the bacteria in
our nostrils) and on the skin.
 This also explains how S. aureus can grow in some
foods with high osmotic pressure (such as ham and
other cured meats) or in low moisture foods that
tend to inhibit the growth of other organisms.
 The yellow pigment probably confers some
protection from the antimicrobial effects of sunlight.
 S. aureus produces many toxins that contribute to
the bacterium’s pathogenicity by increasing its
ability to invade the body or damage tissue.
 The infection of surgical wounds by S. aureus is a
common problem in hospitals. And its ability to
develop resistance quickly to such antibiotics as
penicillin contributes to its danger to patients in
hospital environments.
 S. aureus produces the toxin responsible for toxic
shock syndrome, a severe infection characterized by
high fever and vomiting, some7mes even death.
 S. aureus also produces an enterotoxin that causes
vomiting and nausea when ingested; it is one of the
most common causes of food poisoning.
Order Lactobacillales
 Several important genera are found in the order
Lactobacillales.
 The genus Lactobacillus is a representative of the
industrially important lactic acid–producing bacteria.
 Most lack a cytochrome system and are unable to
use oxygen as an electron acceptor. Unlike most
obligate anaerobes, though, they are aerotolerant
and capable of growth in the presence of oxygen.
But compared to oxygen-utilizing microbes, they
grow poorly. However, the production of lactic acid
from simple carbohydrates inhibits the growth of
competing organisms and allows them to grow
competitively despite their inefficient metabolism.
 The genus Streptococcus shares the metabolic
characteristics of the genus Lactobacillus. There are
several industrially important species, but the
streptococci are best known for their pathogenicity.
 The genera Enterococcus and Listeria are more
conventional metabolically. Both are facultative
anaerobes, and several species are important
pathogens.
Genus Lactobacillus
 The largest genus in this order is Lactobacillus with
around 100 species. Lactobacillus contains non-
sporing rods and some-times coccobacilli that lack
catalase and cytochromes, are usually facultative
anaerobic or microaerophilic, produce lactic acid as
their main or sole fermentation product, and have
complex nutritional requirements.
 Lactobacilli carry out either a homolactic
fermentation using the Embden-Meyerhof pathway
or a heterolac8cfermenta8on with the pentose
phosphate pathway. They grow op8mally under
slightly acidic conditions, when the pH is between
4.5 to 6.4. The genus is found on plant surfaces and
in dairy products, meat, water, sewage, beer, fruits,
and many other materials. Lactobacilli also are part
of the normal flora of the human body in the mouth,
intes8nal tract, and vagina. They usually are not
pathogenic.
 In humans, bacteria of the genus Lactobacillus (lak-
tō-bä-silʹlus) are located in the vagina, intestinal
tract, and oral cavity. Lactobacilli are used
commercially in the production of sauerkraut,
pickles, buttermilk, and yogurt. Typically, a
succession of lactobacilli, each more acid tolerant
 than its predecessor, participates in these lactic acid
fermentations.
 Lactobacillus is indispensable to the food and dairy
industry. Lactobacilli are used in the production of
fermented vegetable foods (sauerkraut, pickles),
beverages (beer, wine, juices), sour dough bread,
Swiss and other hard cheeses, yogurt, and sausage.
Yogurt is probably the most popular fermented milk
product in the United States.
 In commercial production, nonfat or low-fat milk is
pasteurized, cooled to 43°C or lower, inoculated
with Streptococcus thermophilus and Lactobacillus
bulgaricus.
- S. thermophilus grows more rapidly at first and
renders the milk anoxic and weakly acidic.
- L. bulgaricus then acidifies the milk even more.
Acting together, the two species ferment almost all
the lactose to lactic acid and flavor the yogurt with
diacetyl (S. thermophilus) and acetaldehyde (L.
bulgaricus). Fruits or fruit flavors are pasteurized
separately and then combined with the yogurt.
 At least one species, L. plantarum, is sold
commercially as a probiotic agent that may provide
some health benefits for the consumer.
 On the other hand, some lactobacilli also create
problems. They sometimes are responsible for
spoilage of beer, milk, and meat because their
metabolic end products contribute undesirable
flavors and odors.
Genus Streptococcus
 Members of the genus Streptococcus (strep-tō-
kokʹkus) are spherical, gram-positive bacteria that
typically appear in chains. They are a taxonomically
complex group, probably responsible for more
illnesses and causing a greater variety of diseases
than any other group of bacteria.
 Pathogenic streptococci produce several
extracellular substances that contribute to their
pathogenicity. Among them are products that
destroy phagocytic cells that ingest them. Enzymes
produced by some streptococci spread infections by
digesting connective tissue of the host, which may
also result in extensive tissue destruction.
 Infections are also allowed to spread from sites of
injury by enzymes that lyse the fibrin (a threadlike
protein) of blood clots.
 A few nonpathogenic species of streptococci are
important in the production of dairy products
 Grouped into 2 based on hemolysis activity: Beta
and non-beta hemolysis.
 Beta-hemolytic streptococci. A useful basis for the
classification of some streptococci is their colonial
appearance when grown on blood agar. The beta-
hemolytic species produce a hemolysin that forms a
clear zone of hemolysis on blood agar. This group
includes the principal pathogen of the streptococci,
Streptococcus pyogenes (pī-äjʹen-ēz), also known as
the beta-hemolytic group A streptococcus.
- Group A represents one of an antigenic group (A
through G) within the hemolytic streptococci.
Among the diseases caused by S. pyogenes are
scarlet fever, pharyngitis (sore throat), erysipelas,
impetigo, and rheumatic fever.
- The most important virulence factor is the M protein
on the bacterial surface by which the bacteria avoid
phagocytosis.
- Another member of the beta-hemolytic streptococci
is Streptococcus agalactiae (āʹgal-acīē-ī), in the beta-
hemolytic group B. It is the only species with the
group B antigen and is the cause of an important
disease of the newborn, neonatal sepsis.
 Non-beta-hemolytic streptococci. Certain
streptococci are not beta-hemolytic, but when
grown on blood agar, their colonies are surrounded
by a distinctive greening. These are the alpha-
hemolytic streptococci. The greening represents a
partial destruction of the red blood cells caused
mostly by the action of bacteria-produced hydrogen
peroxide, but it appears only when the bacteria
grow in the presence of oxygen. The most important
pathogen in this group is Streptococcus
pneumoniae, the cause of pneumococcal
pneumonia.
 Also included among the alpha-hemolytic
streptococci are species of streptococci called
viridans streptococci.
- However, not all species form the alpha-hemoly8c
greening (virescent = green), so this is not really a
sa8sfactory group name. Probably the most
significant pathogen of the group is Streptococcus
mutans (mūʹtans), the primary cause of dental
caries.
SEE FIGURE 2
Genus Enterococcus and Genus Listeria
 Enterococcus. The enterococci are adapted to areas
of the body that are rich in nutrients but low in
oxygen, such as the gastrointestinal tract, vagina,
and oral cavity. They are also found in large
numbers in human stool.
- Because they are relatively hardy microbes, they
persist as contaminants in a hospital environment,
on hands, bedding, and even as a fecal aerosol. In
recent years they have become a leading cause of
nosocomial infections, especially because of their
high resistance to most antibiotics.
- Two species, Enterococcus faecalis (en-te-rō-kokʹ
kusfe-kāʹ lis) and Enterococcus faecium (fēʹsēum),
are responsible for much of the infec,ons of surgical
wounds and the urinary tract. In medical settings
they frequently enter the bloodstream through
invasive procedures, such as indwelling catheters.
 Listeria. The pathogenic species of the genus
Listeria, Listeria monocytogenes (lis-teʹrē-ämo-nō-sī-
toʹjenēz), can contaminate food, especially dairy
products.
- Important characteristics of L. monocytogenes are
that it survives within phagocytic cells and is capable
of growth at refrigeration temperatures.
- If it infects a pregnant woman, the organism poses
the threat of stillbirth or serious damage to the
fetus.
Genus Leuconostoc
 From family Leuconostocaceae, contains facultative
gram-positive cocci, which may be elongated or
elliptical and arranged in pairs or chains.
 Leuconostocs lack catalase and cytochromes and
carry out heterolactic fermentation by converting
glucose to D-lactate and ethanol or acetic acid by
means of the phosphoketolase pathway.
 They can be isolated from plants, silage, and milk.
The genus is used in wine production, in the
fermentation of vegetables such as cabbage and
cucumbers (pickles), and in the manufacture of
buttermilk, butter, and cheese. L. mesenteroides
synthesizes dextrans from sucrose and is important
in industrial dextran produc0on.
 Leuconostoc species are involved in food spoilage
and tolerate high sugar concentra0ons so well that
they grow in syrup and are a major problem in sugar
refineries.
 Enterococcaceae (Enterococcus) and
Streptococcaceae (Streptococcus, Lactococcus) are
important families of chemo-heterotrophic,
mesophilic, non-sporing, gram-positive cocci. In
practice, they are often distinguished primarily
based on phenotypic properties such as oxygen
relationships, cell arrangement, the presence of
catalase and cytochromes, and peptidoglycan
structure.
SEE FIGURE 3 AND 4
Genus Mycoplasma
 From Order Mycoplasmatales
 The mycoplasmas are highly pleomorphic because
they lack a cell wall and can produce filaments that
resemble fungi, hence their name (mykes= fungus,
and plasma = formed).
 Cells of the genus Mycoplasma (mī-ko-plazʹma) are
very small, ranging in size from 0.1 to 0.25 μm, with
a cell volume that is only about 5% of that of a
typical bacillus.
 Because their size and plasticity allowed them to
pass through filters that retained bacteria, they
were originally considered to be viruses.
Mycoplasmas may represent the smallest self-
replicating organisms that are capable of a free-
living existence.
 One species has only 517 genes; the minimum
necessary is between 265 and 350. Studies of their
DNA suggest that they are genetically related to the
gram-positive bacterial group that includes the
genera Bacillus, Streptococcus, and Lactobacillus but
have gradually lost genetic material. The term
degenerative evolution has been used to describe
this process.
 The most significant human pathogen among the
mycoplasmas is M. pneumoniae (nu-mōʹnē-ī), which
is the cause of a common form of mild pneumonia.
Other genera in the order Mycoplasma-tales are
Spiroplasma (spī-rō-plazʹmä), cells with a tight
corkscrew morphology that are serious plant
pathogens and common parasites of plant-feeding
insects, and Ureaplasma (ū-rē-äplazʹmä), so named
because they can enzymatically hydrolyze the urea
in urine and are occasionally associated with urinary
tract infections.
 Mycoplasmas can be grown on artificial media that
provide them with sterols (if necessary) and other
special nutritional or physical requirements.
Colonies are less than 1 mm in diameter and have a
characteristic “fried egg” appearance when viewed
under magnification.
 For many purposes, cell culture methods are often
more satisfactory. In fact, mycoplasmas grow so well
by this method that they are a frequent
contamination problem in cell culture laboratories.
SEE FIGURE 5
B. ACTINOBACTERIA: The high G+C Gram-positives
Preview
 Bergey’s Manual classifies the actinomycetes and
other high G + C gram-positive bacteria using 16S
rRNA data.
 They are placed in the phylum Actinobacteria, which
is a large, complex grouping with one class and six
orders.
 Actinomycetes are gram-positive, aerobic bacteria,
but are distinctive because they have filamentous
hyphae that differentiate to produce asexual spores.
Many closely resemble fungi in overall morphology.
 The morphology and arrangement of spores, cell
wall chemistry, and the types of sugars present in
cell extracts are particularly important in
actinomycete taxonomy and are used to divide
these bacteria into different groups.
 Actinomycetes have considerable practical impact
because they play a major role in the mineralization
of organic matter in the soil and are the primary
source of most naturally synthesized antibiotics.
 The genera Corynebacterium, Mycobacterium, and
Nocardia include important human pathogens.
 High G + C gram-positive bacteria are in the phylum
Actinobacteria. Many bacteria in this phylum are
highly pleomorphic in their morphology; the genera
Corynebacterium and Gardnerella, for example, and
several genera such as Streptomyces grow only as
extended, often branching filaments.
 Several important pathogenic genera are found in
the Actinobacteria, such as the Mycobacterium
species causing tuberculosis and leprosy. The genera
Streptomyces, Frankia, Actinomyces, and Nocardia
are often informally called a radiate, or starlike,
form of growth by reason of their often-branching
filaments.
 Their morphology resembles that of filamentous
fungi; however, the actinomycetes are prokaryotic
cells, and their filaments have a diameter much
smaller than that of the eukaryotic molds.
- Some actinomycetes further resemble molds by
their possession of externally carried asexual spores
that are used for reproduction.
 Filamentous bacteria, like filamentous fungi, are
very common inhabitants in soil, where a
filamentous pattern of growth has advantages.
 The filamentous organism can bridge water-free
gaps between soil particles to move to a new
nutritional site.
 This morphology also gives the organism a much
higher surface-to-volume ratio and improves its
ability to absorb nutrients in the highly competitive
soil environment.
GENERAL PROPERTIES OF THE ACTINOMYCETES
 Actinomycetes are a fascinating group of
microorganisms.
 source of most of the antibiotics used in medicine
to-day.
 produce metabolites that are used as anticancer
drugs, anti-helminthics (for instance ivermectin,
which is given to dogs to prevent heart worm), and
drugs that suppress the immune system in patients
who have received organ transplants. This practical
aspect of the actinomycetes is linked very closely to
their mode of growth.
 Actinomycetes also have great ecological
significance.
 They are primarily soil inhabitants and are very
widely distributed.
 They can degrade an enormous number and variety
of organic compounds and are extremely important
in the mineralization of organic matter.
 Although most actinomycetes are free-living
microorganisms, a few are pathogens of humans,
other animals, and some plants.
 Like the myxobacteria, the prosthecate bacteria, and
several other microbes like the actinomycetes
undergo a complex life cycle.
 The life cycle of many actinomycetes includes the
development of filamentous cells, called hyphae,
and spores.
 When growing on a solid substratum such as soil or
agar, the actinomycetes develop a branching
network of hyphae. The hyphae grow both on the
surface of the substratum and into it to form a
dense mat of hyphae termed a substrate mycelium.
 Septae usually divide the hyphae into long cells (20
µm and longer) containing several nucleoids. In
many actinomycetes, substrate hyphae differentiate
into upwardly growing hyphae to form an aerial
mycelium that extends above the substratum.
 Because the physiology of the actinomycete has
switched from actively growing vegetative cells into
this special cell type, these compounds are often
called secondary metabolites.
 The aerial hyphae form thin-walled spores upon
septation. These spores are considered exospores
because they do not develop within a mother cell
like the endospores of Bacillus and Clostridium. If
the spores are located in a sporangium, they may be
called sporangiospores.
 The spores can vary greatly in shape. Like spore
formation in other bacteria, actinomycete
sporulation is usually in response to nutrient
deprivation.
 Most actinomycete spores are not particularly heat
resistant but withstand desiccation well, so they
have considerable adaptive value.
 Most actinomycetes are not motile, and spores are
dispersed by wind or adhering to animals; in this
way they may find a new habitat that will provide
needed nutrients. In the few motile genera, motility
is confined to flagellated spores.
FIGURE 6 AND 7
 Actinomycete cell wall composition varies greatly
among different groups and is of considerable
taxonomic importance.
 Four major cell wall types can be distinguished
according to three features of peptidoglycan
composition and structure: the amino acid in
tetrapeptide side chain position 3, the presence of
glycine in interpeptide bridges, and peptidoglycan
sugar content.
 Whole cell extracts of actinomycetes with wall types
II, III, and IV also contain characteristic sugars that
are useful in identification.
 Some other taxonomically valuable properties are:
- the morphology and color of the mycelium and
sporangia
- the surface features and arrangement of spores, the
percent G + C in DNA
- the phospholipid composition of cell membranes,
and spore heat resistance.
 Of course, 16S rRNA sequences has proven valuable.
Several actinomycete genomes have been
sequenced including:
- Mycobacterium tuberculosis
- M. leprae
- Streptomyces coelicolor
- S. avermitilis
SEE FIGURE 8
 It has been clear for some time that some of the
phenotypic traits traditionally used to determine
actinomycete taxonomy do not always fit with 16S
rRNA sequence data.
 Phylogenetic analyses based on 16S rRNA sequences
are used to classify the high G + C gram-positive
bacteria (i.e., those gram-positive bacteria with a
DNA base composition above approximately 50 mol
% G + C).
 The phylum is large and very complex; it contains
one class (Actinobacteria), five sub-classes, six
orders, 14 suborders, and 44 families. • In this
system, the actinobacteria are composed of the
actinomycetes and their high G + C relatives.
Genus Mycobacterium
 The mycobacteria are aerobic, non-endospore-
forming rods.
 The name myco, meaning funguslike, was derived
from their occasional exhibition of filamentous
growth.
 Many of the characteristics of mycobacteria, such as
acid-fast staining, drug resistance, and
pathogenicity, are related to their distinctive cell
wall, which is structurally similar to gram-negative
bacteria.
 The outermost lipopolysaccharide layer in
mycobacteria is replaced by mycolic acids, which
form a waxy, water-resistant layer. This makes the
bacteria resistant to stresses such as drying. Also,
few antimicrobial drugs are able to enter the cell.
 Nutrients enter the cell through this layer very
slowly, which is a factor in the slow growth rate of
mycobacteria; it sometimes takes weeks for visible
colonies to appear. The mycobacteria include the
important pathogens Mycobacterium tuberculosis
(mī-kōbak-tiʹrē-um tü-ber-kū-lōʹsis), which causes
tuberculosis and M. leprae (lepʹrī), which causes
leprosy.
 The mycobacteria are generally separated into two
groups: (1) the slow growers, such as M.
tuberculosis, and (2) the fast, or rapid, growers,
which form visible colonies on appropriate media
within 7 days.
- The slow-growing mycobacteria are more likely to
be pathogenic to humans.
- The rapidly growing group also contains a number of
occasional, nontuberculous human pathogens,
which most commonly infect wounds.
- These mycobacteria are more likely to be
nonpathogenic soil and water microbes.
Genus Corynebacterium
 The corynebacterial (coryne= club-shaped) tend to
be pleomorphic, and their morphology often varies
with the age of the cells.
 The family Corynebacteriaceae has one genus,
Corynebacterium, which contains aerobic and
facultative, catalase-positive, straight to slightly
curved rods, often with tapered ends.
- Club-shaped forms are also seen.
- The bacteria often remain partially attached after
snapping division, resulting in angular arrangements
of the cells, or a palisade arrangement in which rows
of cells are lined up side by side.
 Corynebacteria form metachromatic granules, and
their walls have meso-diaminopimelic acid.
- Although some species are harmless saprophytes,
many corynebacteria are plant or animal pathogens.
 The best-known species is Corynebacterium
diphtheriae (kôrʹi-nēbak-,-rē-um dif-thiʹrē-ī), the
causative agent of diphtheria.
Genus Propionibacterium
 The name of the genus Propionibacterium (prō-pē-
onʹē-bakG-rē-um) is derived from the organism’s
ability to form propionic acid; some species are
important in the fermentation of Swiss cheese.
Propionibacterium acnes (akʹnēz) are bacteria that
are commonly found on human skin and are
implicated as the primary bacterial cause of acne.
Genus Frankia
 The genus Frankia (frankʹē-ä) forms nonmotile
spores in a sporogenous body.
 It grows in symbiotic association with the roots of at
least eight families of higher non-leguminous plants
(e.g., alder trees) and is a microaerophile that can fix
atmospheric nitrogen.
 The roots of infected plants develop nodules that fix
nitrogen so efficiently that a plant such as an alder
can grow in the absence of combined nitrogen (e.g.,
NO3-) when nodulated.
 Within the nodule cells, Frankia forms branching
hyphae with globular vesicles at their ends. These
vesicles may be the sites of nitrogen fixa8on.
 The nitrogen-fixation process resembles that of
Rhizobium in that it is oxygen sensitive and requires
molybdenum and cobalt.
Genus Streptomyces
 The genus Streptomyces (strep-tōmīʹses) is the best
known of the actinomycetes and is one of the
bacteria most commonly isolated from soil.
 The reproductive asexual spores of Streptomyces
are formed at the ends of aerial filaments. If each
spore lands on a suitable substrate, it is capable of
germinating into a new colony.
 These organisms are strict aerobes. They often
produce extracellular enzymes that enable them to
u7lize proteins, polysaccharides (such as starch and
cellulose), and many other organic materials found
in soil.
 Streptomyces characteristically produce a gaseous
compound called geosmin, which gives fresh soil its
typical musty odor.
 Species of Streptomyces are valuable because they
produce most of our commercial antibiotics. This
has led to intensive study of the genus—there are
nearly 500 described species.
Genus Actinomyces
 The genus Actinomyces (ak-Gn-ōmīʹsēs) consists of
facultative anaerobes that are found in the mouth
and throat of humans and animals. They
occasionally form filaments that can fragment.
 One species, Actinomyces israelii (isrāʹlē-ē), causes
actinomycosis, a tissue-destroying disease usually
affecting the head, neck, or lungs.
Genus Nocardia
 The genus Nocardia (nō-kärʹdē-ä) morphologically
resembles Actinomyces; however, these bacteria are
aerobic.
 To reproduce, they form rudimentary filaments,
which fragment into short rods.
 The structure of their cell wall resembles that of the
mycobacteria; therefore, they are often acid-fast.
 Nocardia species are common in soil. Some species,
such as Nocardia asteroids (asʹter-oidēz),
occasionally cause a chronic, difficult-to treat
pulmonary infection.
 N. asteroidesis also one of the causative agents of
mycetoma, a localized destructive infection of the
feet or hands.
Genus Micrococcus
 The genus Micrococcus contains aerobic, catalase-
positive cocci that occur mainly in pairs, tetrads, or
irregular clusters and are usually nonmotile.
 Unlike many other actinomycetes, Micrococcus does
not undergo morphological differentiation.
 Micrococci colonies often are yellow, orange, or red.
They are wide-spread in soil, water, and on
mammalian skin, which may be their normal habitat.
Genus Arthrobacter
 The genus Arthrobacter contains aerobic, catalase
positive rods with respiratory metabolism and lysine
in its peptidoglycan.
 Its most distinctive feature is a rod-coccus growth
cycle.
 When Arthrobacter grows in exponential phase, the
bacteria are irregular, branched rods that may
reproduce by a process called snapping division.
 As they enter stationary phase, the cells change to a
coccoid form. Upon transfer to fresh medium, the
coccoid cells differentiate to form actively growing
rods.
 Although arthrobacters often are isolated from fish,
sewage, and plant surfaces, their most important
habitat is the soil, where they constitute a significant
component of the microbial flora.
 They are well adapted to this niche because they are
very resistant to desiccation and nutrient
deprivation. This genus is unusually flexible
nutritionally and can even degrade some herbicides
and pesticides.
Genus Bifidobacterium
 Bifidobacterium probably is the best-studied genus.
 Bifidobacteria are nonmotile, non-sporing, gram-
positive rods of varied shapes that are slightly
curved and clubbed; often they are branched.
 The rods can be single or in clusters and V-shaped
pairs.
 photosystems I and II and carrying out oxygenic
Bifidobacterium is anaerobic and actively ferments
carbohydrates to produce acetic and lactic acids, but
no carbon dioxide.
 It is found in the mouth and intestinal tract of warm-
blooded vertebrates, in sewage, and in insects.
 B. bifidum is a pioneer colonizer of the human
intestinal tract, particularly when babies are breast
fed.
A few Bifidobacterium infections have been
reported in humans, but the genus does not appear
to be a major cause of disease.
Genus Gardnerella
 Gardnerella vaginalis (gard-ne-relʹla va-jin-alʹis) is a
bacterium that causes one of the most common
forms of vaginitis. There has always been some
difficulty in assigning a taxonomic position in this
species, which is gram-variable, and which exhibits a
highly pleomorphic morphology.
SEE FIGURE 9
II. The Non Proteobacteria Gram-Negative
Bacteria
Preview
 Some bacterial groups, such as those represented by
the hyperthermophiles Aquifex and Thermotoga, are
deeply branching and very old; other bacterial taxa
have arisen more recently.
 All photosynthetic bacteria, including the
cyanobacteria, were once considered a phonetically
unified group.
 Phylogenetic analysis reveals that the purple
photosynthetic bacteria are proteobacteria,
separating from them green sulfur and green non-
sulfur bacteria.
 The cyanobacteria are separated from other
photosynthetic bacteria because they resemble
eucaryotic phototrophs in having both
photosynthesis.
 Their rRNA sequences also indicate that they are
different from other photosynthetic bacteria.
 Bacteria, such as the chlamydiae, that are obligate
intracellular parasites have relinquished some of
their metabolic independence through loss of
metabolic pathway genes. They use their host’s
energy supply and/or cell constituents.
 Gliding motility is widely distributed among bacteria
and is very useful to organisms that digest insoluble
nutrients or move over the surfaces of moist, solid
substrata.
SEE FIGURE 10
Phylum Aquificae and Thermotogae
 Thermophilic microbes are found in both the
bacterial and archaeal domains, but to date, all
hyperthermophilic procaryotes (those with optimum
growth temperatures above 85°C) belong to the
Archaea.
 The phyla Aquificiae and Thermotoga are two
examples of bacterial thermophiles.
 The phylum Aquificae is thought to represent the
deepest or oldest branch of Bacteria. It contains one
class, one order, and eight genera.
 Two of the best-studied genera are Aquifex and
Hydrogenobacter.
 Aquifex pyrophilusis a gram-negative,
microaerophilic rod. It is thermophilic with a
temperature optimum of 85°C and a maximum of
95°C. ‘
 Aquifex is a chemolithoautotroph that captures
energy by oxidizing hydrogen, thiosulfate, and sulfur
with oxygen as the terminal electron acceptor.
 Because Aquifex and Hydrogenobacter are both
thermophilic chemolithoautotrophs, it has been
suggested that the original bacterial ancestor was
probably thermophilic and chemolithoautotrophic.
Phylum Deinococcus-Thermus
 the genus Deinococcus is best studied.
 Deinococci are spherical or rod-shaped with
distinctively different 16S rRNA.
 They often are associated in pairs or tetrads and are
aerobic, mesophilic, and catalase positive; usually
they can produce acid from only a few sugars.
Although they stain gram positive, their cell wall is
layered with an outer membrane like the gram-
negative bacteria.
 They also differ from gram-positive cocci in having L-
ornithine in their peptidoglycan, lacking teichoic
acid, and having a plasma membrane with large
amounts of palmitoleic acid rather than
phosphatidylglycerol phospholipids.
 Almost all strains are extraordinarily resistant to
both desiccation and radiaton; they can survive as
much as 3 to 5 million rad of radiation (an exposure
of 100 rad can be lethal to humans).
Phylum Chlorobi
 The phylum Chlorobi has only one class (Chlorobia),
order (Chlorobiales), and family (Chlorobiaceae).
 The green sulfur bacteria are a small group of
obligately anaerobic photolithoautotrophs that use
hydrogen sulfide, elemental sulfur, and hydrogen as
electron sources.
 sulfide oxidation is deposited outside the cell. Their
photosynthetic pigments are located in ellipsoidal
vesicles called chlorosomes or chlorobium vesicles,
which are attached to the plasma membrane but are
not continuous with it.
- Chlorosomes are the most efficient light-harvesting
complexes found in nature. The chlorosome
membrane is not a normal lipid bilayer. Instead
bacteriochlorophyll molecules are grouped into
lateral arrays held together by carotenoids and
lipids.
- Chlorosomes contain accessory bacteriochlorophyll
pigments, but the reaction center bacterio-
chlorophyll is located in the plasma membrane
where it obtains energy from chlorosome pigments.
These bacteria flourish in the anoxic, sulfide-rich
zones of lakes.
- Although they lack flagella and are nonmotile, some
species have gas vesicles to adjust their depth for
optimal light and hydrogen sulfide. Those forms
without vesicles are found in sulfide-rich muds at
the bottom of lakes and ponds.
 The green sulfur bacteria are very diverse
morphologically. They may be rods, cocci, or vibrios;
some grow singly, and others form chains and
clusters. They are either grass-green or chocolate-
brown in color. Representative genera are
Chlorobium, Prosthecochloris, and Pelodictyon.
Phylum Chloroflexi
 The phylum Chloroflexi has both photosynthetic and
non-photosynthetic members.
 Chloroflexus is the major representative of the
photosynthetic green non-sulfur bacteria. However,
the term “green non-sulfur” is a misnomer because
not all members of this group are green, and some
use sulfur.
 Chloroflexus is a filamentous, gliding, thermophilic
bacterium that often is isolated from neutral to
alkaline hot springs where it grows in the form of
orange-reddish mats, usually in association with
cyanobacteria. It resembles the green sulfur bacteria
with small chlorosomes and accessory
bacteriochlorophyll c.
 Like the purple bacteria, its light-harvesting
complexes contain bacteriochlorophyll a and are in
the plasma membrane.
 Its metabolism is more similar to that of the purple
non-sulfur bacteria. Chloroflexus can carry out
anoxygenic photosynthesis with organic compounds
as carbon sources or grow aerobically as a
chemoheterotroph. It doesn’t appear closely related
 to any other bacterial group based on 16S rRNA
studies and is a deep and ancient branch of the
bacterial tree. Genomic analysis of Chloroflexus
aurantiacus should help elucidate the origin of
photosynthesis.
 The non-photosynthetic, gliding, rod-shaped or
filamentous bacterium Herpetosiphon also is
included in this phylum. Herpetosiphon is an aerobic
chemoorganotroph with respiratory metabolism
that uses oxygen as the electron acceptor. It can be
isolated from freshwater and soil habitats.
Phylum Cyanobacteria
 The cyanobacteria are the largest and most diverse
group of photosynthetic bacteria.
- There is little agreement about the number of
cyanobacterial species. Older classifications had as
many as 2,000 or more species. In one recent
system this has been reduced to 62 species and 24
genera.
 Bergey’s Manual of Systematic Bacteriology
describes 56 genera. Cyanobacterial diversity is
reflected in the G + C content of the group, which
ranges from 35 to 71%.
 Although cyanobacteria are gram-nega0ve bacteria,
their photosynthe0c system closely resembles that
of the eucaryotes because they have chlorophyll a
and photosystems I and II, thereby performing
oxygenic photosynthesis.
 Indeed, the cyanobacteria were once known as
“blue-green algae.” Like the red algae,
cyanobacteria use phycobiliproteins as accessory
pigments. Photosynthetic pigments and electron
transport chain components are located in thylakoid
membranes lined with particles called
phycobilisomes.
 These contain phycobilin pigments, particularly
phycocyanin and phycoerythrin, that transfer energy
to photosystem II. Carbon dioxide is assimilated
through the Calvin cycle; the enzymes needed for
this process are localized to internal structures
called carboxysomes.
 The reserve carbohydrate is glycogen. Sometimes
they store extra nitrogen as polymers of arginine or
aspartic acid in cyanophycin granules. Phosphate is
stored in polyphosphate granules.
- Because cyanobacteria lack the enzyme ⍺-
ketoglutarate dehydrogenase, they do not have a
fully functional citric acid cycle.
 The pentose phosphate pathway plays a central role
in their carbohydrate metabolism. Although many
cyanobacteria are obligate photolithoautotrophs, a
few can grow slowly in the dark as
chemoheterotrophs by oxidizing glucose and a few
other sugars.
 Under anoxic conditions Oscillatoria limnetica
oxidizes hydrogen sulfide instead of water and
carries out anoxygenic photosynthesis much like the
green photosynthetic bacteria. Obviously,
cyanobacteria are capable of considerable metabolic
flexibility.
 Cyanobacteria also vary greatly in shape and
appearance. They range in diameter from about 1 to
10 µm and may be unicellular, exist as colonies of
many shapes, or form filaments called trichomes.
 A trichome is a row of bacterial cells that are in close
contact with one another over a large area.
Although many appear blue-green because of
phycocyanin, isolates from the open ocean are red
or brown in color because of the pigment
phycoerythrin.
 Cyanobacteria modulate the relative amounts of
these pigments in a process known as chromatic
adaptation.
- When orange light is perceived, phycocyanin
production is stimulated, while blue and blue-green
light promote the production of phycoerythrin. It is
thought that sensory rhodopsins may play a role in
signaling the spectral quality of light.
 Many cyanobacterial species use gas vacuoles to
position themselves in optimum illumination in the
water column— a form of phototaxis.
 Gliding motility is used by other cyanobacteria.
Although cyanobacteria lack flagella, about one-
third of the marine Synechococcus strains swim at
rates up to 25 µm/sec by an unknown mechanism.
- Swimming motility is not used for phototaxis;
instead, it appears to be used in chemotaxis toward
simple nitrogenous compounds such as urea.
 Cyanobacteria show great diversity with respect to
reproduction and employ a variety of mechanisms:
binary fission, budding, fragmentation, and multiple
fission.
 In the last process a cell enlarges and then divides
several times to produce many smaller progenies,
which are released upon the rupture of the parental
cell. Fragmentation of filamentous cyanobacteria
can generate small, motile filaments called
hormogonia.
 Some species develop akinetes, specialized,
dormant, thick-walled resting cells that are resistant
to desiccation. Often these germinate to form new
filaments.
 Many filamentous cyanobacteria fix atmospheric
nitrogen by means of special cells called heterocysts.
- Around 5 to 10% of the cells develop into
heterocysts when these cyanobacteria are deprived
of both nitrate and ammonia, their preferred
nitrogen sources. When individual cyanobacterial
cells in a filamentous chain differentiate into
heterocysts, the heterocysts develop a very thick cell
wall.
- Within these specialized cells, photosynthetic
membranes are reorganized and the proteins that
make up photosystem II and phycobilisomes are
degraded.
 Photosystem I remains functional to produce ATP,
but no oxygen is generated. This inability to
generate O2 is critical because the enzyme
nitrogenase is extremely oxygen sensitive.
 The thick heterocyst wall slows or prevents O2
diffusion into the cell, and any O2 present is
consumed during respiration.
 Heterocyst structure and physiology ensure that it
remains anaerobic; it is dedicated to nitrogen
fixation and does not replicate.
 It obtains nutrients from adjacent vegetative cells
and contributes fixed nitrogen in the form of amino
acids. Nitrogen fixation also is carried out by some
cyanobacteria that lack heterocysts. Some fix
nitrogen under dark, anoxic conditions in microbial
mats.
 Planktonic forms such as Trichodesmium fix nitrogen
and contribute significantly to the marine nitrogen
budget.
SEE FIGURE 11
Phylum Plantomycetes
 The phylum Planctomycetes contains one class, one
order, and four genera. The planctomycetes are
morphologically unique bacteria, having
compartmentalized cells.
 Although each species is unique, all follow a basic
cellular organization that includes a cytoplasmic
membrane closely surrounded by the cell wall,
which lacks peptidoglycan.
 The largest internal compartment, the
intracytoplasmic membrane (ICM), is separated
from the cytoplasmic membrane by a peripheral,
ribosome-free region called the paryphoplasm.
- The nucleoid of the planctomycete Gemmata
obscuriglobus is located in the nuclear body, which
is enclosed in a double membrane. The species
“Candidatus Brocadia anammoxidans” does not
localize its DNA within a nuclear body; however, it
has another compartment: the anammoxosome.
- This is the site of anaerobic ammonia oxidation, a
unique and recently discovered form of
chemolithotrophyin which ammonium ion (NH4+)
serves as the electron donor a nitrite (NO2-) as the
terminal electron acceptor; it is reduced to nitrogen
gas (N2). Note that the nomenclature of this
microbe is still in flux so in some cases genus and
species names are enclosed in quotation marks.
Biogeochemical cycling: Nitrogen cycle.
 The genus Planctomyces attaches to surfaces
through a stalk and holdfast; the other genera in the
order lack stalks.
 Most of these bacteria have life cycles in which
sessile cells bud to produce motile swarmer cells.
The swarmer cells are flagellated and swim for a
while before settling down to attach and begin
reproduction.
Phylum Chlamydiae
 The gram-negative Chlamydiae are obligate
intracellular parasites. That is, they must grow and
reproduce within host cells.
- Although their ability to cause disease is widely
recognized, many species grow within protists and
animal cells without adverse affects. It is thought
that these hosts represent a natural reservoir for the
Chlamydiae.
 The phylum Chlamydiae has one class, one order,
four families, and only six genera.
 The genus Chlamydia is by far the most important
and best studied; it will be the focus of our
attention.
 Chlamydiae are nonmotile, coccoid bacteria, ranging
in size from 0.2 to 1.5 µm. They reproduce only
within cytoplasmic vesicles of host cells by a unique
developmental cycle in-volving the formation of two
cell types: elementary bodies and reticulate bodies.
- Although their envelope resembles that of other
gram-negative bacteria, the cell wall differs in
lacking muramic acid and a peptidoglycan layer.
- Elementary bodies achieve osmotic stability by
cross-linking their outer membrane proteins, and
possibly periplasmic proteins, with disulfide bonds.
 Chlamydiae are extremely limited metabolically,
relying on their host cells for key metabolites. This is
reflected in the size of their genome. It is relatively
small at 1.0 to 1.3 Mb; the G + C content is 41 to
44%.
- Chlamydial reproduction begins with the attachment
of an elementary body (EB) to the cell surface.
- Elementary bodies are 0.2 to 0.6 µm in diameter,
contain electron-dense nuclear material and a rigid
cell wall, and are infectious.
- The host cell phagocytoses the EB, which are held in
inclusion bodies where the EB reorganizes itself to
form a reticulate body (RB) or initial body. The RB is
specialized for reproduction rather than infection.
Reticulate bodies are 0.6 to 1.5 µm in diameter and
have less dense nuclear material and more
ribosomes than EBs; their walls are also more
flexible.
- About 8 to 10 hours after infection, the reticulate
body undergoes binary fission and RB reproduction
continues until the host cell dies. A chlamydia-filled
inclusion can become large enough to be seen in a
light microscope and even fill the host cytoplasm.
- After 20 to 25 hours, RBs begin to differentiate into
infectious EBs and continue this process until the
host cell lyses and releases the chlamydiae 48 to 72
hours after infection.
FIGURE 12
 Chlamydial metabolism is very different from that of
other gram-negative bacteria. It had been thought
that chlamydiae cannot catabolize carbohydrates or
other substances or synthesize ATP.
 Chlamydia psittaci, one of the best-studied species,
lacks both flavoprotein and cytochrome electron
transport chain carriers, but has a membrane
translocase that acquires host ATP in exchange for
ADP.
- Chlamydiae seem to be energy parasites that are
completely dependent on their hosts for ATP.
However, this might not be the complete story. The
C. trachomatis genome sequence indicates that the
bacterium may be able to synthesize at least some
ATP.
- Although there are two genes for ATP/ADP
translocases, there also are genes for substrate-level
phosphorylation, electron transport, and oxiditive
phosphorylation.
 When supplied with precursors from the host, RBs
can synthesize DNA, RNA, glycogen, lipids, and
proteins.
- The RBs have porins and active membrane transport
proteins, but little is known about these. They also
can synthesize at least some amino acids and
coenzymes. The EBs have minimal metabolic ac0vity
and cannot take in ATP or synthesize proteins. They
are designed exclusively for transmission and
infection. Insights from microbial genomes:
Genomic analysis of pathogenic microbes.
 Three chlamydial species are important pathogens
of hu-mans and other warm-blooded animals. C.
trachomatis infects humans and mice. In humans it
causes trachoma, nongonococcal urethritis, and
other diseases. C. psittaci causes psittacosis in
humans.
- Unlike C. trachoma8s, it also infects many other
animals (e.g., parrots, turkeys, sheep, caBle, and
cats) and invades the intes0nal, respiratory, and
genital tracts; the placenta and fetus; the eye; and
the synovial fluid of joints.
- Chlamydiophila pneumoniae is a common cause of
human pneumonia.
SEE FIGURE 13
Phylum Spirochaetes
 Phylum Spirochaetes (Greek spira, a coil, and
chaete, hair) contains gram-negative,
chemoheterotrophic bacteria distinguished by their
structure and mechanism of motility.
 They are slender, long bacteria (0.1 to 3.0 µm by 5
to 250 µm) with a flexible, helical shape. Many
species are so slim that they are only clearly visible
in a light microscope by means of phase-contrast or
dark-field optics.
 Spirochetes differ greatly from other bacteria with
respect to motility and can move through very
viscous solutions though they lack external rotating
flagella.
- When in contact with a solid surface, they exhibit
creeping or crawling movements. Their unique
pattern of motility is due to an unusual
morphological structure called the axial filament.
 The distinctive features of spirochete morphology
are evident in electron micrographs.
 The central protoplasmic cylinder contains
cytoplasm and the nucleoid, and is bounded by a
plasma membrane and a gram-negative cell wall.
 Two to more than a hundred flagella, called axial
fibrils, periplasmic flagella, or endoflagella, extend
from both ends of the cylinder and often overlap
one another in the center third of the cell.
- The whole complex of periplasmic flagella, the axial
filament, lies inside a flexible outer sheath.
- The outer sheath contains lipid, protein, and
carbohydrate and varies in structure between
different genera.
- Its precise function is unknown, but the sheath is
essential because spirochetes die if it is damaged or
removed.
 The outer sheath of Treponema pallidum has few
proteins exposed on its surface. This allows the
syphilis spirochete to avoid attack by host
antibodies.
 The way in which periplasmic flagella propel the cell
has not been fully established. Mutants with straight
rather than curved flagella are nonmotile.
 The periplasmic flagella rotate like the external
flagella of other bacteria.
 This causes the corkscrew-shaped outer sheath to
rotate and move the cell through the surrounding
liquid.
 Flagellar rotation may also flex or bend the cell and
account for the crawling movement seen on solid
surfaces.
 Spirochetes can be anaerobic, facultatively
anaerobic, or aerobic.
 Carbohydrates, amino acids, long-chain fatty acids,
and long-chain fatty alcohols may serve as carbon
and energy sources.
 The group is exceptionally diverse ecologically and
grows in habitats ranging from mud to the human
mouth.
 Members of the genus Spirochaeta are free-living
and often grow in anoxic and sulfide-rich freshwater
and marine environments. Some species of the
genus Leptospira grow in toxic water and moist soil.
 Some spirochetes form symbiotic associations with
other organisms and are found in a variety of
locations: the hindguts of termites and wood-eating
roaches, the digestive tracts of mollusks (Cristispira)
and mammals, and the oral cavities of animals
(Treponema denticola, T. oralis).
 Spirochetes from termite hindguts and freshwater
sediments have nitrogenase and can fix nitrogen.
 There is evidence that they contribute significantly
to the nitrogen nutrition of termites. Spirochetes
coat the surfaces of many protozoa from termite
and wood-eating roach hindguts.
 For example, the flagellate Myxotricha paradoxais
covered with slender spirochetes (0.15 by 10 µm in
 length) that are firmly attached and help move the
protozoan.
 Some members of the genera Treponema, Borrelia,
and Leptospira are important pathogens; for
example, Treponema pallidum causes syphilis, and
Borrelia burgdorferi is responsible for Lyme disease.
 The study of T. pallidum and its role in syphilis has
been hindered by the inability to culture the
spirochete outside its human host. The T. pallidum
genome sequence shows that this spirochete is
metabolically crippled and quite dependent on its
host.
 The B. burgdorferi genome consists of a linear
chromosome of 910,725 base pairs and at least 17
linear and circular plasmids, which constitute
another 533,000 base pairs.
 The plasmids have some genes that are normally
found on chromosomes, and plasmid proteins seem
to be involved in bacterial virulence.
SEE FIGURE 14
Phylum Bacteriodetes
 The phylum Bacteroidetes is very diverse and seems
most closely related to the phylum Chlorobi. The
phylum has three classes (Bacteroides,
Flavobacteria, and Sphingobacteria), 12 families, and
63 genera.
 The class Bacteroides contains anaerobic, gram-
negative, non-sporing, motile or nonmotile rods of
various shapes.
 These bacteria are chemoheterotrophic and usually
produce a mixture of organic acids as fermentation
end products. They do not reduce sulfate or other
sulfur compounds.
- The genera are identified using properties such as
general shape, motility and flagellation pattern, and
fermentation end products. These bacteria grow in
habitats such as the oral cavity and intestinal tract of
vertebrates and the rumen of ruminants.
- Although difficulty culturing these anaerobes has
hindered our understanding of them; they are
clearly widespread and important. Often they
benefit their host. Bacteroides ruminicolais a major
component of the rumen flora; it ferments starch,
pectin, and other carbohydrates.
 About 30% of the bacteria isolated from human
feces are members of the genus Bacteroides, and
these organisms may provide extra nutrition by
degrading cellulose, pectins, and other complex
carbohydrates. The family also is involved in human
disease.
 Members of the genus Bacteroides are associated
with diseases of major organ systems, ranging from
the central nervous system to the skeletal system.
 B. fragilis is a particularly common anaerobic
pathogen found in abdominal, pelvic, pulmonary,
and blood infections.
 Another important group in the Bacteroidetes is the
class Sphingobacteria. Besides the similarity in their
16S rRNA sequences, sphingobacteria often have
sphingolipids in their cell walls. Some genera in this
class are Sphingobacterium, Saprospira, Flexibacter,
Cytophaga, Sporocytophaga, and Crenothrix.
 The genera Cytophaga, Sporocytophaga, and
Flexibacter differ from each other in morphology,
life cycle, and physiology.
 Bacteria of the genus Cytophaga are slender rods,
often with pointed ends. Sporocytophaga is similar
to Cytophaga but forms spherical resting cells called
microcysts. Flexibacter produces long, flexible
threadlike cells when young and is unable to use
complex polysaccharides. Often colonies of these
bacteria are yellow to orange because of carotenoid
or flexirubin pigments.
 Some of the flexirubins are chlorinated, which is
unusual for biological molecules.
 Members of the genera Cytophaga and
Sporocytophaga are aerobes that actively degrade
complex polysaccharides. Soil cy-tophagas digest
cellulose; both soil and marine forms attack chitin,
pectin, and keratin.
 Some marine species even degrade agar, a
component of seaweed. Cytophagas play a major
role in the mineralization of organic matter and can
cause great damage to exposed fishing gear and
wooden structures. They also are a major
component of the bacterial population in sewage
treatment plants and presumably contribute
significantly to the waste treatment process.
Wastewater treatment.
 Although most cytophaga sare free-living, some can
be isolated from vertebrate hosts and are
pathogenic.
 Cytophaga columnaris and others cause diseases
such as columnaris disease, cold water disease, and
fin rot in freshwater and marine fish.
 The gliding motility so characteristic of these
organisms is quite different from flagellar motility.
- Gliding motility is present in a wide diversity of taxa:
fruiting and nonfruiting aerobic chemoheterotrophs,
cyanobacteria, green nonsulfur bacteria, and at least
two gram-positive genera (Heliobacterium and
Desulfonema).
- Gliding bacteria lack flagella and are stationary while
suspended in liquid medium. When in contact with a
surface, they glide along, leaving a slime trail.
Movement can be very rapid; some cytophagas
travel 150µm in a minute, whereas filamentous
gliding bacteria may reach speeds of more than
600µm/minute.
 Young organisms are the most motile, and motility
often is lost with age. Low nutrient levels usually
stimulate gliding. The gliding mechanism is not well
understood.
 Gliding motility gives a bacterium many advantages.
Many aerobic chemoheterotrophic gliding bacteria
actively digest insoluble macromolecular substrates
 such as cellulose and chitin, and gliding mo0lity is
ideal for searching these out.
 Because many of the digestive enzymes are cell wall
associated, the bacteria must be in contact with
insoluble nutrient sources; gliding motility makes
this possible.
 Gliding movement is well adapted to drier habitats
and to movement within solid masses such as soil,
sediments, and rotting wood that are permeated by
small channels. G
 Gliding bacteria, like flagellated bacteria, can
position themselves at optimal conditions of light
intensity, oxygen, hydrogen sulfide, temperature,
and other factors that influence growth and survival.
FIGURE 1

FIGURE 2
FIGURE 3

FIGURE 4.1
FIGURE 4.2

FIGURE 5

FIGURE 6
FIGURE 7

FIGURE 8
FIGURE 9.1

FIGURE 9.2
FIGURE 10.1

FIGURE 10.2
FIGURE 11

FIGURE 12
FIGURE 13

FIGURE 14