MICTOTAXONOMY 03
photosynthetic ancestor.
Domain Bacteria: The Proteobacteria
Phylum Protobacteria
 The phylum Proteobacteria is the largest,
phylogenetically coherent bacterial group with over
2,000 species assigned to more than 500 genera.
 Many of these gram-negative bacteria are of
considerable importance, either as disease agents or
because of their contributions to ecosystems.
- The bacteria such as Escherichia coli, are major
experimental organisms studied in many
laboratories.
 These bacteria are very diverse in their metabolism
and lifestyles, which range from obligate
intracellular parasitism to a free-living existence in
soil and aquatic habitats.
 Chemolithotrophic bacteria obtain energy and
electrons by oxidizing inorganic compounds rather
than the organic nutrients employed by many
bacteria.
- They often have substantial ecological impact
because of their ability to oxidize many forms of
inorganic nitrogen and sulfur.
 Some Proteobacteria produce specialized structures
such as prosthecae, stalks, buds, sheaths, or
complex fruiting bodies.
 Many bacteria that specialize in predatory or
parasitic modes of existence, such as Bdellovibrio
and the rickettsias, have relinquished some of their
metabolic independence through the loss of
metabolic pathways.
- They depend on the prey’s or host’s energy supply
and/or cell constituents.
Did you know?
 The proteobacteria, which includes most of the
gram-negative, chemoheterotrophic bacteria, are
presumed to have arisen from a common
important bacteria capable of inducing nitrogen
 They are now the largest taxonomic group of
bacteria. However, few are now photosynthetic;
other metabolic and nutritional capacities have
arisen to replace this characteristic.
 The phylogenetic relationship in these groups is
based upon rRNA studies.
- The name Proteobacteria was taken from the
mythological Greek god Proteus, who could assume
many shapes.
 The proteobacteria are separated into five classes
designated by Greek letters.
A. Class Alphaproteobacteria
 As a group, the ⍺-proteobacteria includes most of
the proteo-bacteria that are capable of growth at
very low levels of nutrients.
- Some have unusual morphology, including
protrusions such as stalks or buds known as
prosthecae.
 The alphaproteobacteria also include agriculturally
fixation in symbiosis with plants, and several plant
and human pathogens.
 Some have unusual metabolic modes such as
methylotrophy (Methylobacterium),
chemolithotrophy (Nitrobacter), and the ability to fix
nitrogen (Rhizobium).
 Members of genera such as Rickettsia and Brucella
are important pathogens; in fact, Rickettsia is an
obligate intracellular parasite.
The Purple Non-Sulfur Bacteria
 All the purple bacteria use anoxygenic
photosynthesis, possess bacteriochlorophylls a or b,
and have their photosynthetic apparatus in
membrane systems that are continuous with the
plasma membrane.
 Most are motile by polar flagella. All purple
nonsulfur bacteria are ⍺-proteobacteria, except for
Rhodocyclus (β-proteobacteria).
 The purple nonsulfur bacteria are exceptionally
flexible in their choice of an energy source.
- Normally they grow anaerobically as
photoorganoheterotrophs; they trap light energy
and employ organic molecules as both electron and
carbon sources.
 Although they are called nonsulfur bacteria, some
species can oxidize very low, nontoxic levels of
sulfide to sulfate, but they do not oxidize elemental
sulfur to sulfate.
 In the absence of light, most purple nonsulfur
bacteria can grow aerobically as
chemoorganoheterotrophs, but some species carry
out fermentations anaerobically.
- Oxygen inhibits bacteriochlorophyll and carotenoid
synthesis so that cultures growing aerobically in the
dark are colorless.
 Purple nonsulfur bacteria vary considerably in
morphology.
- They may be spirals (Rhodospirillum), rods
(Rhodopseudomonas), half circles or circles
(Rhodocyclus), or they may even form prosthecae
and buds (Rhodomicrobium).
- Because of their metabolism, they are most
prevalent in the mud and water of lakes and ponds
with abundant organic matter and low sulfide levels.
- There also are marine species.
 Rhodospirillum and Azospirillum (both in the family
Rhodospirillaceae) are among several bacterial
genera capable of forming cysts.
- These resting cells differ from the well-characterized
endospores made by gram-positive bacteria such as
Bacillus and Clostridium.
- Cyst-forming bacteria are not limited to ⍺-
proteobacteria; for instance, Azotobacter, a β-
proteobacterium, also forms cysts.
Genus Pelagibacter
 One of the most abundant microorganisms on Earth,
certainly in the ocean environment, is Pelagibacter
ubique (pel-ajʹē-bak-teruʹbēk).
 It is a member of a group of marine microbes
discovered by use of the FISH (Fluorescence in situ
hybridization) technique and named SAR 11 because
of their original discovery in the Sargasso Sea. P.
ubique is the first member of this group to be
successfully cultivated.
o Its genome has been sequenced and found to have
only 1354 genes.
o This number is very low for a free-living organism,
although several mycoplasmas have even fewer
genes.
o Bacteria in a symbiotic relationship have lower
metabolic requirements and have the smallest
genomes.
o The bacterium is extremely small, a little over 0.3
μm diameter.
o Its small size and minimal genome probably give it a
competitive advantage for survival in low-nutrient
environments.
o In fact, it seems to be the most abundant living
organism (part of its name, ubique, is derived from
ubiquitous), on the basis of weight, in the oceans,
where its sheer numbers must give it an important
role in the Earth’s carbon cycle.
Genus Ehrlichia
 Ehrlichiae are gram-negative, rickettsia-like bacteria
that live obligately within white blood cells.
 Ehrlichia (ėrʹlik-ē-ä) species are transmitted by ticks
to humans and cause ehrlichiosis, a sometimes-fatal
disease.
Genus Azospirillum
 Agricultural microbiologists have been interested in
members of the genus Azospirillum (ā-zōspīʹril-lum),
a soil bacterium that grows in close association with
the roots of many plants, especially tropical grasses.
 It uses nutrients excreted by the plants and in return
fixes nitrogen from the atmosphere.
 This form of nitrogen fixation is most significant in
some tropical grasses and in sugar cane, although
the organism can be isolated from the root system
of many temperate-climate plants, such as corn.
 The prefix azo-is frequently encountered in
nitrogen-fixing genera of bacteria. It is derived from
a (without) and zo (life), about the early days of
chemistry when oxygen was removed, by a burning
candle, from an experimental atmosphere.
Genus Rickettsia
 In earlier editions of Bergey’s Manual, the genera
Rickettsia, Coxiella, and Chlamydia were grouped
closely because they share the common
characteristic of being obligate intracellular
parasites—that is, they reproduce only within a
mammalian cell.
- In the second edition they are now widely
separated.
 Rickettsias are gram-negative rod-shaped bacteria,
or coccobacilli.
 One distinguishing feature of most rickettsias is that
they are transmitted to humans by bites of insects
and ticks, unlike the Coxiella (discussed later with 𝛾-
proteobacteria).
 Rickettsia enter their host cell by inducing
phagocytosis.
 They quickly enter the cytoplasm of the cell and
begin reproducing by binary fission.
 They can usually be cultivated artificially in cell
culture or chick embryos.
 The rickettsias are responsible for a number of
diseases known as the spotted fever group.
- These include epidemic typhus, caused by Rickettsia
prowazekii (ri-ketʹsē-ä prou-wä-zeʹkēē) and
transmitted by lice.
- endemic murine typhus caused by R. typhi (tiʹfē) and
transmitted by rat fleas.
- and Rocky Mountain spotted fever, caused by R.
ricke/siiri-ketʹsē-ē) and transmitted by ticks.
 In humans, rickettsial infections damage the
permeability of blood capillaries, which results in a
characteristic spotted rash.
Genus Rhizobium, Bradyrhizobium, and Agrobacterium
 The Rhizobium (rī-zōʹbē-um) and Bradyrhizobium
(brād-ē-rī-zōʹbē-um) are two of the more important
genera of a group of agriculturally important
bacteria that specifically infect the roots of
leguminous plants, such as beans, peas, or clover.
- For simplicity these bacteria are known by the
common name of rhizobia.
- The presence of rhizobia in the roots leads to
formation of nodules in which the rhizobia and plant
form a symbiotic relationship, resulting in the
fixation of nitrogen from the air for use by the plant.
 Like rhizobia, the genus Agrobacterium (agʹrō-bak-
tʹrē-um) can invade plants.
- These bacteria do not induce root nodules or fix
nitrogen.
- Of particular interest is Agrobacterium tumefaciens.
This is a plant pathogen that causes a disease called
crown gall; the crown is the area of the plant where
the roots and stem merge. The tumorlike gall is
induced when A. tumefaciens inserts a plasmid
containing bacterial genetic information into the
plant’s chromosomal DNA.
Genus Acetobacter and Gluconobacter
 Acetobacter (äʹsē-tō-bak-tėr) and Gluconobacter
(glüʹkonō-bak-tėr) are industrially important aerobic
organisms that convert ethanol into acetic acid
(vinegar).
Genus Wolbachia
 Wolbachia (wol-baʹkē-ä) are probably the most
common infectious bacterial genus in the world.
 Even so, little is known about Wolbachia; they live
only inside the cells of their hosts, usually insects (a
relationship known as endosymbiosis).
- These intracellular bacteria infect mainly arthropod
species, including a high proportion of insects, and
some nematodes.
- It is one of the most common parasitic microbes and
is possibly the most common reproductive parasite
in the biosphere.
 Therefore, Wolbachia escape detection by the usual
culture methods.
Genus Nitrobacter and Nitrosomonas
 Nitrobacter (nī-trō-bakʹtėr) and Nitrosomonas (nī-
trō-sō-mōʹnas) are genera of nitrifying bacteria that
are of great importance to the environment and to
agriculture.
 They are chemoautotrophs capable of using
inorganic chemicals as energy sources and carbon
dioxide as the only source of carbon, from which
they synthesize all of their complex chemical
makeup.
 The energy sources of the genera Nitrobacter and
Nitrosomonas (the latter is a member of the
betaproteobacteria) are reduced nitrogenous
compounds.
 Nitrobacter species oxidize ammonium (NH4+) to
nitrite (NO2−), which is in turn oxidized by
Nitrosomonas species to nitrates (NO3−) in the
process of nitrification.
 Nitrate is important to agriculture; it is a nitrogen
form that is highly mobile in soil and therefore likely
to be encountered and used by plants.
SEE FIGURE 1
Genus Caulobacter and Hyphomicrobium
 Members of the genus Caulobacter (kô-lō-bakʹtėr)
are found in low-nutrient aquatic environments,
such as lakes.
- They feature stalks that anchor the organisms to
surfaces.
- This arrangement increases their nutrient uptake
because they are exposed to a continuously
changing flow of water and because the stalk
increases the surface-to-volume ratio of the cell.
- When the nutrient concentration is exceptionally
low, the size of the stalk increases, evidently to
provide an even greater surface area for nutrient
absorption.
 Budding bacteria do not divide by binary fission into
two nearly identical cells.
- The budding process resembles the asexual
reproductive processes of many yeasts.
 The parent cell retains its identity while the bud
increases in size until it separates as a completely
new cell.
 An example is the genus Hyphomicrobium (hī-fō-mī-
krōʹbē-um).
 These bacteria, like the caulobacteria, are found in
low-nutrient aquatic environments and have even
been found growing in laboratory water baths.
 Both Caulobacter and Hyphomicrobium produce
prominent prosthecae.
Genus Brucella and Bartonella
 Brucella (brüʹsel-la) bacteria are small nonmotile
coccobacilli.
- All species of Brucella are obligate parasites of
mammals and cause the disease brucellosis.
- Its medical interest is the ability of Brucella to
survive phagocytosis, an important element of the
body’s defense against bacteria.
 The genus Bartonella (bärʹtō-nel-la) contains several
members that are human pathogens.
 The best known is Bartonella henselae, a gram-
negative bacillus that causes cat-scratch disease.
B. Class Betaproteobacteria
 The β-proteobacteria overlap the ⍺-proteobacteria
metabolically but tend to use substances that diffuse
from organic decomposition in the anoxic zone of
habitats.
 Some of these bacteria use hydrogen, ammonia,
methane, volatile fatty acids, and similar substances.
 As with the ⍺-proteobacteria, there is considerable
metabolic diversity; the β-proteobacteria may be
chemoheterotrophs, photolithotrophs,
methylotrophs, and chemolithotrophs.
 The class Betaproteobacteria has seven orders and
12 families.
 Several important pathogenic bacteria are found in
this group.
Genus Thiobacillus
 From Order Hydrogenophilales
- Bergey’s Manual disperses these bacteria between
two classes; for example, Thiobacillus and
Macromonas are β-proteobacteria, whereas
Thiomicrospira, Thiobacterium, Thiospira, Thiothrix,
Beggiatoa, and others are 𝛾-proteobacteria.
- Only some of these bacteria have been isolated and
studied in pure culture.
- Most is known about the genera Thiobacillus and
Thiomicrospira.
- Thiobacillus is a gram-negative rod, and
Thiomicrospira is a long spiral cell; both have polar
flagella. They differ from many of the nitrifying
bacteria in that they lack extensive internal
membrane systems.
 Thiobacillus (thī-ō-bä-silʹlus) species and other
sulfur-oxidizing bacteria are important in the sulfur
cycle.
 These chemoautotrophic bacteria are capable of
obtaining energy by oxidizing the reduced forms of
sulfur, such as hydrogen sulfide (H2S), or elemental
sulfur (S0), into sulfates (SO42−).
 one of the best-studied chemolithotrophs and most
prominent of the colorless sulfur bacteria. Like the
nitrifying bacteria, colorless sulfur bacteria are a
highly diverse group.
 Many are unicellular rod-shaped or spiral sulfur-
oxidizing bacteria that are nonmotile or motile by
flagella.
SEE FIGURE 2
Genus Spirillium
 The habitat of the genus Spirillum (spī-rilʹlum) is
mainly fresh water. An important morphological
difference from the helical spirochetes is that
Spirillum bacteria are motile by conventional polar
flagella, rather than axial filaments.
 The spirilla are relatively large, gram-negative,
aerobic bacteria.
 Spirillum volutans (vō-lū-tans) is often used as a
demonstration slide when microbiology students are
first introduced to the operation of the microscope.
Genus Sphaerotilus
 Sheathed bacteria, which include Sphaerotilus
natans (sfe-räʹ8-lusnaʹtans), are found in freshwater
and in sewage.
 These gram-negative bacteria with polar flagella
form a hollow, filamentous sheath in which to live.
 Sheaths are protective and aid in nutrient
accumulation.
 Sphaerotilus probably contributes to bulking, an
important problem in sewage treatment.
Genus Burkholderia
 The genus Burkholderia was formerly grouped with
the genus Pseudomonas, which is now classified
under the gamma proteobacteria. •
 Like the pseudomonads, almost all Burkholderia
species are motile by a single polar flagellum or tuft
of flagella.
- The best-known species is the aerobic, gram-
negative rod Burkholderia cepacia (berkʹhōld-ėr-ē-
äse-pāʹse-ä).
 It has an extraordinary nutritional spectrum and is
capable of degrading more than 100 different
organic molecules.
 This capability is often a factor in the contamination
of equipment and drugs in hospitals; these bacteria
may actually grow in disinfectant solutions.
 This bacterium is also a problem for persons with
the genetic lung disease cystic fibrosis, in whom it
metabolizes accumulated respiratory secretions.
 Burkholderia pseudomallei (sūdo-malʹ lē-ī) is a
resident in moist soils and is the cause of a severe
disease (melioidosis) endemic in southeast Asia and
northern Australia.
Genus Neisseria, Bordetella, and Zoogloea
 Bacteria of the genus Neisseria (nī-seʹrē-ä) are
aerobic, gram-negative cocci that usually inhabit the
mucous membranes of mammals.
- Pathogenic species include the gonococcus
bacterium Neisseria gonorrhoeae (go-nôr-rēʹī), the
causative agent of gonorrhoea and N. meningitidis
(men-nin-jiʹG-dis), the agent of meningococcal
meningitis.
 Of special importance is the nonmotile, aerobic,
gram-negative rod Bordetella pertussis (bôrʹde-tel-
läpėr-tusʹsis).
- This serious pathogen is the cause of pertussis, or
whooping cough.
 The genus Zoogloea (zōʹō-glē-ä) is important in the
context of aerobic sewage-treatment processes,
such as the activated sludge system.
 As they grow, Zoogloea bacteria form fluffy, slimy
masses that are essential to the proper operation of
such systems.
C. Class Gammaproteobacteria
 The 𝛾-proteobacteria constitute the largest
subgroup of the proteobacteria and include a great
variety of physiological types.
 Many important genera are chemoorganotrophic
and facultatively anaerobic. Other genera contain
aerobic chemoorganotrophs, photolithotrophs,
chemolithotrophs, or methylotrophs.
 According to some DNA-rRNA hybridization studies,
the 𝛾-proteobacteria are composed of several
deeply branching groups.
- One consists of the purple sulfur bacteria; a second
includes the intracellular parasites Legionella and
Coxiella.
 The two largest groups contain a wide variety of
non-photosynthetic genera. Ribosomal RNA
superfamily I is represented by the families
Vibrionaceae, Enterobacteriaceae, and
Pasteurellaceae.
- These bacteria use the Embden-Meyerhof and
pentose phosphate pathways to catabolize
carbohydrates. Most are facultative anaerobes.
- Ribosomal RNA superfamily II contains mostly
aerobes that often use the Entner-Doudoroffand
pentose phosphate pathways to catabolize many
different kinds of organic molecules.
- The genera Pseudomonas, Azotobacter, Moraxella,
Xanthomonas, and Acinetobacter belong to this
superfamily.
 The exceptional diversity of these bacteria is evident
from the fact that Bergey’sManual divides the class
Gammaproteobacteria into 14 orders and 28
families.
SEE FIGURE 3
Genus Beggiatoa
 Beggiatoa alba (bejʹjē-ä-tō-äalʹba), the only species
of this unusual genus, grows in aquatic sediments at
the interface between the aerobic and anaerobic
layers.
 Morphologically, it resembles certain filamentous
cyanobacteria, but it is not photosynthetic.
 Motility is by gliding. The mechanism is the
production of slime, which attaches to the surface
on which movement occurs and provides lubrication
allowing the organism to glide.
 Nutritionally B. alba uses hydrogen sulfide (H2S) as
an energy source and accumulates internal granules
of sulfur.
 The ability of this organism to obtain energy from an such as soap residues or cap liner adhesives found in
inorganic compound was an important factor in the a solution, has been unexpectedly troublesome.
discovery of autotrophic metabolism.  Pseudomonads are even capable of growth in some
antiseptics, such as quaternary ammonium
Genus Francisella
compounds.
 Francisella (franʹsis-elʹlä) is a genus of small, - Their resistance to most antibiotics has also been a
pleomorphic bacteria that grow only on complex source of medical concern.
media enriched with blood or tissue extracts. - This resistance is probably related to the
 Francisella tularensis (tüʹlär-en-sis) causes the characteristics of the cell wall porins, which control
disease tularemia. the entrance of molecules through the cell wall.
- The large genome of pseudomonads also codes for
Genus Pseudomonas several very efficient efflux pump systems that eject
antibiotics from the cell before they can function.
 Members of the order Pseudomonadales are gram-
Pseudomonads are responsible for about one in ten
negative aerobic rods or cocci. The most important
nosocomial infections (hospital-acquired infections),
genus in this group is Pseudomonas (sū-dō-mōʹnas)
especially among infections in burn units.
consists of aerobic, gram-negative rods that are
 Persons with cystic fibrosis are also especially prone
motile by polar flagella, either single or tufts.
to infections by Pseudomonas and the closely
 Pseudomonads are very common in soil and other
related Burkholderia.
natural environments.
 The pseudomonads have a great practical impact in
 Many species of pseudomonads excrete
several ways, including these:
extracellular, water-soluble pigments that diffuse
1. Many can degrade an exceptionally wide variety of
into their media.
organic molecules.
- One species, Pseudomonas aeruginosa (ā-rü-ji-
- Thus, they are very important in the mineralization
nōʹsä), produces a soluble, blue-green pigmentation.
process (the microbial breakdown of organic
Under certain conditions, particularly in weakened
materials to inorganic substances) in nature and in
hosts, this organism can infect the urinary tract,
sewage treatment.
burns, and wounds, and can cause blood infections,
- The fluorescent pseudomonads can use
abscesses, and meningitis.
approximately 80 different substances as their
- Other pseudomonads produce soluble fluorescent
carbon and energy sources.
pigments that glow when illuminated by ultraviolet
2. Several species (e.g., P. aeruginosa) are important
light. One species, P. syringae (sėrʹin-gī), is an
experimental subjects.
occasional plant pathogen. (Some species of
- Many advances in microbial physiology and
Pseudomonas have been transferred, based upon
biochemistry have come from their study. For
rRNA studies, to the genus Burkholderia, which was
example, the study of P. aeruginosa has significantly
discussed previously with the betaproteobacteria.)
advanced our understanding of how bacteria form
 In hospitals and other places where pharmaceutical
biofilms and the role of extracellular signaling in
agents are prepared, the ability of pseudomonads to
bacterial communities and pathogenesis.
grow on minute traces of unusual carbon sources,
- The genome of P. aeruginosa has an unusually large
number of genes for catabolism, nutrient transport,
the efflux of organic molecules, and metabolic
regulation. This may explain its ability to grow in
many environments and resist antibiotics.
3. Some pseudomonads are major animal and plant
pathogens. P. aeruginosa infects people with low
resistance such as cystic fibrosis patients.
4. It also invades burns and causes urinary tract
infections. P. syringaeis an important plant
pathogen.
5. Pseudomonads such as P. fluorescens are involved
in the spoilage of refrigerated milk, meat, eggs, and
seafood because they grow at 4°C and degrade
lipids and proteins.
Genus Azotobacter and Azomonas
 Some nitrogen-fixing bacteria, such as Azotobacter
(ā-zō-tō-bakʹtėr) and Azomonas (ā-zō-mōʹnas), are
free-living in soil. These large, ovoid, heavily
capsulated bacteria are frequently used in
laboratory demonstrations of nitrogen fixation.
However, to fix agriculturally significant amounts of
nitrogen, they would require energy sources, such
as carbohydrates, that are in limited supply in soil.
 The genus Azotobacter also is in the family
Pseudomonadaceae. The genus contains large,
ovoid bacteria, 1.5 to 2.0 m in diameter, that may be
motile by peritrichous flagella. The cells are often
pleomorphic, ranging from rods to coccoid shapes,
and form cysts as the culture ages. The genus is
aerobic, catalase positive, and fixes nitrogen non-
symbiotically. Azotobacter is widespread in soil and
water.
Genus Moraxella
 Members of the genus Moraxella (mô-raks-elʹlä) are
strictly aerobic coccobacilli—that is, intermediate in
shape between cocci and rods. Moraxella lacunata
(la-kü-näʹtä) is implicated in conjunctivitis, an
inflammation of the conjunctiva, the membrane that
covers the eye and lines the eyelids.
Genus Acinetobacter
 The genus Acinetobacter (a-si-nēʹtō-bak-tėr) is
aerobic and in stained preparations typically forms
pairs.
 The bacteria occur naturally in soil and water. A
member of this genus, Acinetobacter baumanii
(bouʹman-ē-ē), is an increasing concern to the
medical community because of the rapidity with
which it becomes resistant to antibiotics. Some
strains are resistant to most available antibiotics.
 Not yet widespread in the United States, A.
baumaniiis an opportunistic pathogen primarily
found in a hospital setting. The antibiotic resistance
of the pathogen, combined with the weakened
health of infected hospital patients, has resulted in
an unusually high mortality rate. A. baumaniiis
primarily a respiratory pathogen, but it also infects
skin and soc tissues and wounds and occasionally
invades the bloodstream.
 It is more environmentally hardy than most gram-
negative bacteria, and, once established in a
hospital, it becomes difficult to eliminate.
Order Legionellales
 The genera Legionella and Coxiella are closely
associated in the second edition of Bergey’s Manual,
where both are placed in the same order,
Legionellales.
 Because the Coxiella share an intracellular lifestyle
with the rickettsial bacteria, they were previously
considered rickettsial in nature and grouped with
them. Legionella bacteria grow readily on suitable
artificial media.
Genus Legionella
 Legionella (lē-jä-nelʹlä) bacteria were originally
isolated during a search for the cause of an outbreak
of pneumonia now known as legionellosis.
 The search was difficult because these bacteria did
not grow on the usual laboratory isolation media
then available. After intensive effort, special media
were developed that enabled researchers to isolate
and culture the first Legionella.
 Microbes of this genus are now known to be
relatively common in streams, and they colonize
such habitats as warmwater supply lines in hospitals
and water in the cooling towers of air conditioning
systems. An ability to survive and reproduce within
aquatic amoebas often makes them difficult to
eradicate in water systems.
Genus Coxiella
 Coxiella burnetii (käks-ė-elʹ lä bėr-neʹtē-ē), which
causes Q fever, was formerly grouped with the
rickettsia. Like them, Coxiella bacteria require a
mammalian host cell to reproduce.
 Unlike rickettsias, Coxiella bacteria are not
transmitted among humans by insect or tick bites.
Although cattle ticks harbor the organism, it is most
commonly transmitted by aerosols or contaminated
milk. A spore-like body is present in C. burnetii. This
might explain the bacterium’s relatively high
resistance to the stresses of airborne transmission
and heat treatment.
Order Vibrionales
 Members of the order Vibrionales are facultatively
anaerobic gram-negative rods. Many are slightly
curved. They are found mostly in aquatic habitats.
 There are eight genera in the family: Vibrio,
Photobacterium, Salinivibrio, Listonella, Allomonas,
Enterovibrio, Catencoccus, and Grimontia.
 Several vibrios are important pathogens. Vibrio
cholerae causes cholera, and V. parahaemolyticus
 can cause gastroenteritis in humans following
consumption of contaminated seafood. V.
anguillarum and others are responsible for fish
diseases.
GenusVibrio
 Members of the genus Vibrio (vibʹrē-ō) are rods that
are often slightly curved.
 One important pathogen is Vibrio cholerae (kolʹer-ī),
the causative agent of cholera.
- The disease is characterized by a profuse and watery
diarrhea. V. parahaemolyticus (pa-rä-hē-mō-liʹti-kus)
causes a less serious form of gastroenteritis.
- Usually inhabiting coastal salt waters, it is
transmitted to humans mostly by raw or under
cooked shellfish.
 Several members of the family are unusual in being
bioluminescent. Vibrio fischeri, V. harveyi, and at
least two species of Photobacteriumare among the
few marine bacteria capable of bioluminescence and
emit a blue-green light because of the ac3vity of the
enzyme luciferase.
 The peak emission of light is usually between 472
and 505 nm, but one strain of V. fischeri emits
yellow light with a major peak at 545 nm. Although
many of these bacteria are free-living, V. fischeri, V.
harveyi, P. phosphoreum, and P. leiognathi live
symbiotically in the luminous organs of fish and
squid.
SEE FIGURE 4
Order Enterobacteriales
 The members of the order Enterobacteriales are
facultatively anaerobic, gram-negative rods that are,
if motile, peritrichously flagellated.
- Morphologically, the rods are straight. This is an
important bacterial group, often commonly called
enterics. They inhabit the intestinal tracts of
humans and other animals.
- Most enterics are active fermenters of glucose and
other carbohydrates.
 Enterics have fimbriae that help them adhere to
surfaces or mucous membranes.
- Specialized sex pili aid in the exchange of genetic
information between cells, which often includes
antibiotic resistance.
 Enterics, like many bacteria, produce proteins called
bacteriocins that cause the lysis of closely related
species of bacteria.
- Bacteriocins may help maintain the ecological
balance of various enterics in the intestines.
Genus Escherichia
 The bacterial species Escherichia coli is one of the
most common inhabitants of the human intestinal
tract and is probably the most familiar organism in
microbiology.
 Known about the biochemistry and genetics of E.
coli, and it continues to be an important tool for
basic biological research— many researchers
consider it almost a laboratory pet.
 E. coli is not usually pathogenic.
 However, it can be a cause of urinary tract
infections, and certain strains produce enterotoxins
that cause traveler’s diarrhea and occasionally cause
very serious foodborne disease (E. coli O157:H7).
 Its presence in water or food is an indication of fecal
contamination.
SEE FIGURE 5
Genus Salmonella
 Almost all members of the genus Salmonella
(salʹmön-el-lä) are potentially pathogenic.
- Accordingly, there are extensive biochemical and
serological tests to clinically isolate and identify
salmonellae.
- Salmonellae are common inhabitants of the
intestinal tracts of many animals, especially poultry
and cattle. Under unsanitary conditions, they can
contaminate food.
 The nomenclature of the genus Salmonella is
unusual. Instead of multiple species, members of
the genus Salmonella that are infectious to warm-
blooded animals can be considered for practical
purposes to be a single species, Salmonella enterica
(en-terʹi-kä).
- This species is divided into more than 2400 serovars,
that is, serological varieties. The term serotype is
often used to mean the same thing. By way of
explanation of these terms, when salmonellae are
injected into appropriate animals, their flagella,
capsules, and cell walls serve as antigens that cause
the animals to form antibodies in their blood that
are specific for each of these structures.
 A serovar such as Salmonella typhimurium (m-fi-
murʹē-um) is not a species and should be more
properly written as “Salmonella enterica serovar
Typhimurium.”
 The convention now used by the Centers for Disease
Control and Prevention (CDC) is to spell out the
entire name at the first mention and then
abbreviate it as, for example, Salmonella
Typhimurium. For simplicity, we will identify
serovars of salmonellae in this text as we would
species, that is, S. typhimurium, etc.
Genus Shigella
 Species of Shigella (shi-gelʹlä) are responsible for a
disease called bacillary dysentery, or shigellosis.
 Unlike salmonellae, they are found only in humans.
Some strains of Shigella can cause life-threatening
dysentery.
Genus Klebsiella
 Members of the genus Klebsiella (kleb-sē-elʹlä) are
commonly found in soil or water.
 Many isolates are capable of fixing nitrogen from
the atmosphere, which has been proposed as being
a nutritional advantage in isolated populations with
little protein nitrogen in their diet.
 The species Klebsiella pneumoniae (nü-mōʹnē-ī)
occasionally causes a serious form of pneumonia in
humans.
Genus Serratia
 Serratia marcescens (ser-räʹ tē-ämär-sesʹsens) is a
bacterial species distinguished by its production of
red pigment.
 In hospital situations, the organism can be found on
catheters, in saline irrigation solutions, and in other
supposedly sterile solutions. Such contamination is
probably the cause of many urinary and respiratory
tract infections in hospitals.
Genus Proteus
 Colonies of Proteus (prōʹ tē-us) bacteria growing on
agar exhibit a swarming type of growth.
 Swarmer cells with many flagella move outward on
the edges of the colony and then revert to normal
cells with only a few flagella and reduced motility.
 Periodically, new generations of highly motile
swarmer cells develop, and the process is repeated.
As a result, a Proteus colony has the distinctive
appearance of a series of concentric rings.
 This genus of bacteria is implicated in many
infections of the urinary tract and in wounds.
Genus Yersinia
 Yersinia pestis (yėr-sinʹē-äpesʹFs) causes plague, the
Black Death of medieval Europe.
 Urban rats in some parts of the world and ground
squirrels in the American South-west carry these
bacteria.
 Fleas usually transmit the organisms among animals
and to humans, although contact with respiratory
droplets from infected animals and people can be
involved in transmission.
SEE FIGURE 6
Genus Erwinia
 Erwinia (ėr-wiʹnē-ä) species are primarily plant
pathogens; some cause plant soc-rot diseases.
 These species produce enzymes that hydrolyze the
pectin between individual plant cells. This causes the
plant cells to separate from each other, a disease
that plant pathologists term plant rot.
Genus Enterobacter
 Two Enterobacter (en-te-rō-bakʹtėr) species, E.
cloacae (klō-āʹkī), and E. aerogenes (ā-räʹjen-ēz), can
cause urinary tract infections and hospital-acquired
infections.
 They are widely distributed in humans and animals,
as well as in water, sewage, and soil.
Genus Pasteurella
 The genus Pasteurella (pas-tyėr-elʹ lä) is primarily
known as a pathogen of domestic animals. It causes
sepsis in cattle, fowl cholera in chickens and other
fowl, and pneumonia in several types of animals.
 The best-known species is Pasteurella multocida
(mul-tōʹsi-dä), which can be transmitted to humans
by dog and cat bites.
Genus Haemophilus
 Haemophilus (hē-mäʹ fil-us) is a very important
genus of pathogenic bacteria.
- These organisms inhabit the mucous membranes of
the upper respiratory tract, mouth, vagina, and
intestinal tract.
- The best-known species that affects humans is
Haemophilus influenzae (in-flü-enʹ za), named long
ago because of the erroneous belief that it was
responsible for influenza.
 The name Haemophilus is derived from the
bacteria’s requirement for blood in their culture
medium (hemo= blood).
- They are unable to synthesize important parts of the
cytochrome system needed for respiration, and they
obtain these substances from the heme fraction,
known as the X factor, of blood hemoglobin.
- The culture medium must also supply the
ofactornicotinamide adenine dinucleotide (from
either NAD+ or NADP+), which is known as V factor.
Clinical laboratories use tests for the requirement of
X and V factors to identify isolates as Haemophilus
species.
 Haemophilus influenzae is responsible for several
important diseases. It has been a common cause of
meningitis in young children and is a frequent cause
of earaches.
- Other clinical conditions caused by H. influenzae
include epiglottis (a life-threatening condition in
which the epiglottis becomes infected and
inflamed), septic arthrtis in children, bronchitis, and
pneumonia. Haemophilus ducreyi (dü-krāʹē) is the
cause of the sexually transmitted disease chancroid.
SEE FIGURE 7
D. Class Deltaproteobacteria
 The 𝛅-proteobacteria are distinctive in that they
include some bacteria that are predators on other
bacteria.
 are not a large assemblage of genera, they show
considerable morphological and physiological
 diversity. These bacteria can be divided into two
general groups, all of them chemoorganotrophs.
 Some genera are predators such as the bdellovibrios
and myxobacteria. Others are anaerobes that
generate sulfide from sulfate and sulfur while
oxidizing organic nutrients. The class has eight
orders and 20 families. •
 Bacteria in this group are also important
contributors to the sulfur cycle.
Genus Bdellovibrio
 Bdellovibrio (del-lō-vibʹrē-ō) is a particularly
interesting genus.
 It attacks other gram-negative bacteria. It attaches
tightly (bdella= leech), and after penetrating the
outer layer of gram-negative bacteria, it reproduces
within the periplasm.
 The cell elongates into a tight spiral, which then
fragments almost simultaneously into several
individual flagellated cells. The host cell then lyses,
releasing the Bdellovibrio cells.
Genus Desulfovibrio
 The best-studied sulfur-reducing genus is
Desulfovibrio (dēʹsul-fō-vibʹrē-ō), which is found in
anaerobic sediments and in the intestinal tracts of
humans and animals.
 Sulfur-reducing and sulfate-reducing bacteria use
organic com-pounds such as lactate, ethanol, or
fatty acids as electron donors.
 This reduces sulfur or sulfate to H2S. When H2S
reacts with iron it forms insoluble FeS, which is
responsible for the black color of many sediments.
Genus Myxococcus
 Vegetative cells of the myxobacteria (myxo=nasal
mucus) move by gliding and leave behind a slime
trail.
 Myxococcus xanthus (micks-ō-kokʹkuszanʹthus) and
M. fulvus (fulʹvus) are well-studied representatives
of the genus.
 As they move, their source of nutrition is the
bacteria they encounter, enzymatically lyse, and
digest.
 Large numbers of these gram-negative microbes
eventually aggregate SEE FIGURE 8
 Where the moving cells aggregate, they differen3ate
and form a macroscopic stalked frui3ng body that
contains large numbersofres3ng cells called
myxospores.
 Differentiation is usually triggered by low nutrients.
 Under proper conditions, usually a change in
nutrients, the myxospores germinate and form new
vegetative gliding cells.
E. Class Epsilonproteobacteria
 The 𝜺-proteobacteria slender gram-negative rods
that are helical or curved.
 are the smallest of the five proteobacterial classes.
 They all are slender gram-negative rods, which can
be straight, curved, or helical.
 The 𝜺-proteobacteria have one order,
Campylobacterales, and three families:
Campylobacteraceae, Helicobacteraceae, and the
recently added Nautiliaceae.
 Two pathogenic genera, Campylobacter and
Helicobacter, are microaerophilic, motile, helical or
vibrioid, gram-negative rods.
Genus Campylobacter
 Members of the genus Campylobacter (kamʹpi-lō-
bak-tėr) are microaerophilic vibrios; each cell has
one polar flagellum.
 contains both nonpathogens and species pathogenic
for humans and other animals. ‘
 C. fetus causes reproductive disease and abortions
in cattle and sheep. It is associated with a variety of
conditions in humans ranging from septicemia
(pathogens or their toxins in the blood) to enteritis
(inflammation of the intestinal tract). C. jejunicauses
abortion in sheep and enteritis diarrhea in humans.
Genus Helicobacter
 Members of the genus Helicobacter are micro-
aerophilic curved rods with multiple flagella.
 The species Helicobacter pylori (hēʹlik-ō-bak-tėrpī-
lōrʹē) has been identified as the most common cause
of peptic ulcers in humans and a cause of stomach
cancer.
 There are at least 23 species of Helicobacter, all
isolated from the stomachs and upper intestines of
humans, dogs, cats, and other mammals.
- In developing countries 70 to 90% of the population
is infected; developed countries range from 25 to
50%.
- Most infections are probably acquired during
childhood, but the precise mode of transmission is
unclear. The major human pathogen is Helicobacter
pylori, which causes gastritis and peptic ulcer
disease. H. pylori produces large quantities of
urease, and urea hydrolysis appears to be associated
with its virulence.
SEE FIGURE 9
FIGURE 1

FIGURE 2
FIGURE 3

FIGURE 4
FIGURE 5

FIGURE 6

FIGURE 7
FIGURE 8

FIGURE 9