Domain Bacteria:

the Proteobacteria

MCB100-Module 3

MARK ANTHONY I. JOSE

Microbiology and Biotechnology Group
Department of Biological Sciences
College of Science and Mathema;cs
MSU-Iligan Ins;tute of Technology
Module Outline

I. Phylum Proteobacteria
II. Class Alphaproteobacteria
III. Class Betaproteobacteria
IV. Class Gammaproteobacteria
V. Class Deltaproteobacteria
VI. Class Epsilonproteobacteria
Module Objec/ves

• To be acquainted with the largest phylum in the Domain Bacteria

• To know the different major groups of Phylum Proteobacteria
• To know the major Genera under the different classes of Phylum Proteobacteria.
• To get acquainted with clinically significant members of Proteobacteria.
 Phylum Proteobacteria

• The phylum Proteobacteria is the largest, phylogene2cally coherent

bacterial group with over 2,000 species assigned to more than 500
genera.
• Many of these gram-nega2ve bacteria are of considerable importance,
either as disease agents or because of their contribu2ons 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 parasi2sm to a free-living existence in
soil and aqua2c habitats.
Phylum Proteobacteria

• Chemolithotrophic bacteria obtain energy and electrons by

oxidizing inorganic compounds rather than the organic
nutrients employed by many bacteria.
• They o9en have substan;al 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 frui;ng bodies.
• Many bacteria that specialize in predatory or parasi;c modes
of existence, such as Bdellovibrio and the rickeAsias, 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
cons4tuents.
 • The proteobacteria, which includes most of the
gram-nega4ve, chemoheterotrophic bacteria, are
presumed to have arisen from a common
photosynthe4c ancestor.
• They are now the largest taxonomic group of
bacteria. However, few are now photosynthe4c;
other metabolic and nutri4onal capaci4es have
arisen to replace this characteris4c.
• The phylogene4c rela4onship in these groups is
based upon rRNA studies.
Did you know? • 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 leEers:
• Alphaproteobacterial
• Betaproteobacteria,
• Gammaproteobacteria
• Deltaproteobacteria
• Epsilonproteobacteria.
 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 important bacteria capable of inducing nitrogen
fixaGon 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 Ricke6sia and Brucella are important pathogens; in fact, Ricke6sia 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 photosynthe7c apparatus in membrane systems that are con7nuous with the plasma
membrane.
• Most are mo7le by polar flagella. All purple nonsulfur bacteria are ⍺-proteobacteria, except for
Rhodocyclus (β-proteobacteria).
• The purple nonsulfur bacteria are excep7onally 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 fermenta7ons anaerobically.
• Oxygen inhibits bacteriochlorophyll and carotenoid synthesis so that cultures growing aerobically in the dark
are colorless.
The Purple Non-Sulfur Bacteria

• 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 ma>er 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 res@ng cells differ from the well-characterized endospores made by gram-posi@ve 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-ter uʹbēk).
• It is a member of a group of marine microbes discovered by use of the FISH
(Fluorescence in situ hybridiza7on) 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 cul7vated.
ü Its genome has been sequenced and found to have only 1354 genes.
ü This number is very low for a free- living organism, although several
mycoplasmas have even fewer genes.
ü Bacteria in a symbio7c rela7onship have lower metabolic requirements
and have the smallest genomes.
ü The bacterium is extremely small, a liVle over 0.3 μm diameter.
ü Its small size and minimal genome probably give it a compe77ve
advantage for survival in low-nutrient environments.
ü 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 This Photo by Unknown Author is licensed under CC BY-SA

oceans, where its sheer numbers must give it an important role in the
Earth’s carbon cycle.
• Ehrlichiae are gram-nega2ve,
rickeDsia-like bacteria that live
obligately within white blood
cells.
• Ehrlichia (ėrʹlik-ē-ä) species are
transmiDed by 2cks to humans
and cause ehrlichiosis, a
some2mes fatal disease
This Photo by Unknown Author is licensed under CC BY-SA

Genus Ehrlichia
 Genus Azospirillum
• Agricultural microbiologists have been interested
in members of the genus Azospirillum (ā-zō-
spīʹril-lum), a soil bacterium that grows in close
associaGon 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 fixaGon 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), in reference to the
early days of chemistry when oxygen was Azospirillium Photo by Unknown Author is licensed under CC BY

removed, by a burning candle, from an

experimental atmosphere.
 Genus Ricke6sia
• In earlier edi*ons of Bergey’s Manual, the genera
Ricke3sia, Coxiella, and Chlamydia were grouped closely
be-cause they share the common characteris*c of being
obligate intracellular parasites—that is, they reproduce
only within a mammalian cell.
• In the second ediGon they are now widely separated.
• Ricke>sias are gram-nega*ve rod-shaped bacteria, or
coccobacilli.
• One disGnguishing feature of most rickeSsias is that they are
transmiSed to humans by bites of insects and Gcks, unlike the
Coxiella (discussed later with 𝛾-proteobacteria).
• RickeSsia 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 culGvated arGficially in cell culture or chick
embryos.
 Genus Ricke6sia
• The rickeDsias are responsible for a number of
diseases known as the spoDed fever group.
These include epidemic typhus, caused by
Ricke/sia prowazekii (ri-ketʹsē-ä prou-wä- zeʹkē-
ē) and transmiDed by lice; endemic murine
typhus, caused by R. typhi (Mʹfē) and transmiDed
by rat fleas; and Rocky Mountain spoDed fever,
caused by R. ricke/sii ri-ketʹsē-ē) and transmiDed
by 2cks.
• In humans, rickeDsial infec2ons damage the
permeability of blood capillaries, which results in
a characteris2c spoDed 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 forma*on
of nodules in which the rhizobia and plant form a
symbio*c rela*onship, resul*ng in the fixa*on of
nitrogen from the air for use by the plant.
 Genus Rhizobium, Bradyrhizobium, and Agrobacterium

Like rhizobia, the genus Agrobacterium (agʹrō-bak-7ʹrē-um) has

the ability to invade plants.
These bacteria do not induce root nodules or fix nitrogen.
Of par7cular 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 gene7c
informa7on into the plant’s chromosomal DNA.
Genus Acetobacter and
Gluconobacter • Acetobacter (äʹsē-tō-bak-tėr)
and Gluconobacter (glüʹkon-
ō-bak-tėr) are industrially
important aerobic organisms
that convert ethanol into
aceGc acid (vinegar).

Acetobacter Photo by Unknown Author is licensed under CC BY-SA

Genus Wolbachia
• Wolbachia (wol-baʹkē-ä) are probably the most common infec7ous
bacterial genus in the world.
• Even so, liVle is known about Wolbachia; they live only inside the cells
of their hosts, usually insects (a rela7onship known as endosymbiosis).
• These intracellular bacteria infects mainly arthropod species, including a high
propor;on of insects, and also some nematodes.
• It is one of the most common parasi;c microbes and is possibly the most
common reproduc;ve parasite in the biosphere
Wolbachia Photo
• Therefore, Wolbachia escape detec7on by the usual culture methods.
h"ps://www.worldmosquitoprogram.org/sites/default/files/styles/our_method_image/public/201910/1924%20wolbachi
a.jpg?h=2a479378&itok=XQgWlMky
 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 laDer 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 nitrifica/on.
• 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.
 Genus Caulobacter and
Hyphomicrobium
• Members of the genus Caulobacter (kô-lō-bakʹtėr)
are found in low-nutrient aqua;c 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 con9nuously
changing flow of water and because the stalk
increases the surface-to-volume ra9o of the cell.
• When the nutrient concentra9on is excep9onally
low, the size of the stalk increases, evidently to
provide an even greater surface area for nutrient
absorp9on.

• Budding bacteria do not divide by binary fission into two nearly iden7cal cells.
• The budding process resembles the asexual reproduc7ve processes of many yeasts.
• The parent cell retains its iden7ty while the bud increases in size un7l it separates as a complete new cell.
• An example is the genus Hyphomicrobium (hī-fō-mī-krōʹbē-um).
• These bacteria, like the caulobacteria, are found in low-nutrient aqua7c 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
nonmoJle coccobacilli.
• All species of Brucella are obligate parasites of
mammals and cause the disease brucellosis.
• Its medical interest is the ability of Brucella to
Brucella Photo by Unknown Author is licensed under CC BY-SA

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-
nega2ve bacillus that causes cat-scratch disease. h"ps://i0.wp.com/images-prod.healthline.com/hlcmsresource/images/galleries/cat-scratch-fever/4609-blister-
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 Class Betaproteobacteria

• The β-proteobacteria overlap the ⍺-proteobacteria

metabolically but tend to use substances that diffuse from
organic decomposi8on in the anoxic zone of habitats.
• Some of these bacteria use hydrogen, ammonia, methane,
vola8le fa?y 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-nega4ve 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 nonmo8le or mo8le by
flagella.
 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 moGle by convenGonal polar flagella, rather than axial
filaments.
• The spirilla are relaGvely large, gram-negaGve, aerobic bacteria.
• Spirillum volutans (vō-lū-tans) is o[en used as a demonstraGon slide when microbiology students are first
introduced to the operaGon of the microscope
 Genus Sphaero4lus

• Sheathed bacteria, which include Sphaero7lus natans (sfe-räʹ8-lus naʹtans), are found in freshwater
and in sewage.
• These gram-nega8ve bacteria with polar flagella form a hollow, filamentous sheath in which to live.
• Sheaths are protec8ve and also aid in nutrient accumula8on.
• Sphaero7lus 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 gammaproteobacteria.
• Like the pseudomonads, almost all Burkholderia species are moGle by a
single polar flagellum or tu[ of flagella.
• The best known species is the aerobic, gram-nega@ve rod Burkholderia cepacia
(berkʹhōld-ėr-ē-ä se-pāʹse-ä).
• It has an extraordinary nutriGonal spectrum and is capable of degrading
more than 100 different organic molecules.
• This capability is o[en a factor in the contaminaGon of equipment and
drugs in hospitals; these bacteria may actually grow in disinfectant
soluGons.
• This bacterium is also a problem for persons with the geneGc lung
disease cysGc fibrosis, in whom it metabolizes accumulated respiratory
secreGons.
• 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-nega*ve cocci that usually inhabit the mucous
membranes of mammals.
• Pathogenic species include the gonococcus bacterium Neisseria
gonorrhoeae (go-nôr-rēʹī), the causaGve agent of gonorrhoea and
N. meningiFdis (men-nin-jiʹG-dis), the agent of meningococcal
meningiGs.
• Of special importance is the nonmo*le, aerobic, gram-
nega*ve rod Bordetella pertussis (bôrʹde-tel-lä pėr-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
ac*vated sludge system.
• As they grow, Zoogloea bacteria form fluffy, slimy masses that are
essenGal to the proper operaGon of such systems.
 Class Gammaproteobacteria

• The 𝛾-proteobacteria cons7tute the largest subgroup of the proteobacteria and include a great variety of physiological
types.
• Many important genera are chemoorganotrophic and faculta7vely anaerobic. Other genera contain aerobic
chemoorganotrophs, photolithotrophs, chemolithotrophs, or methylotrophs.
• According to some DNA-rRNA hybridiza7on 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-photosynthe7c 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 faculta=ve anaerobes.
• Ribosomal RNA superfamily II contains mostly aerobes that oCen use the Entner- Doudoroff and pentose phosphate pathways to catabolize many different kinds of organic molecules.
• The genera Pseudomonas, Azotobacter, Moraxella, Xanthomonas, and Acinetobacter belong to this superfamily.

• The excep7onal diversity of these bacteria is evident from the fact that Bergey’s Manual divides the class
Gammaproteobacteria into 14 orders and 28 families.
 Genus Beggiatoa

• Beggiatoa alba (bejʹjē-ä-tō-ä alʹba), the only species of this unusual genus,
grows in aquaJc sediments at the interface between the aerobic and anaerobic
layers.
• Morphologically, it resembles certain filamentous cyanobacteria, but it is not
photosyntheJc.
• MoJlity is by gliding. The mechanism is the producJon of slime, which aDaches
to the surface on which movement occurs and also provides lubricaJon allowing
the organism to glide.
• NutriJonally 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 inorganic compound was
an important factor in the discovery of autotrophic metabolism.
 Genus Francisella

• Francisella (franʹsis-elʹlä) is a genus of

small, pleo- morphic bacteria that
grow only on complex media enriched
with blood or 2ssue extracts.
• Francisella tularensis (tüʹlär-en-sis)
causes the disease tularemia.

Figure 6. Fulfilment of Koch's postulates in Nile (wild type) tilapia. (A) Naïve tilapia fingerlings prior to the
challenge. (B,C) Reproduction of clinical signs and gross lesions post challenge. Inset in (C): streaking of spleen with
widespread multifocal white nodules onto CHAH. (D) Recovery of pure colonies from the spleen of experimentally infected
fish.

Source: Ramírez-Paredes, J. G., Thompson, K. D., Metselaar, M., Shahin, K., Soto, E., Richards, R. H., ... & Adams, A. (2017). A polyphasic
approach for phenotypic and geneKc characterizaKon of the fasKdious aquaKc pathogen Francisella noatunensis subsp.
orientalis. Fron-ers in microbiology, 8, 2324. hQps://doi.org/10.3389/fmicb.2017.02324
 Genus Pseudomonas

• Members of the order Pseudomonadales are gram-negaJve aerobic rods or cocci. The
most important genus in this group is Pseudomonas (sū-dō- mōʹnas) consists of
aerobic, gram-negaJve rods that are moJle by polar flagella, either single or tuXs.
• Pseudomonads are very common in soil and other natural environments.
• Many species of pseudomonads excrete extracellular, water- soluble pigments that
diffuse into their media.
• One species, Pseudomonas aeruginosa (ā-rü-ji-nōʹsä), produces a soluble, blue-green
pigmenta8on. Under certain condi8ons, par8cularly in weakened hosts, this organism can infect
the urinary tract, burns, and wounds, and can cause blood infec8ons, abscesses, and meningi8s.
• Other pseudomonads pro- duce soluble fluorescent pigments that glow when illuminated by
ultraviolet light. One species, P. syringae (sėrʹin-gī), is an occasional plant pathogen. (Some
species of Pseudomonas have been transferred, based upon rRNA studies, to the genus
Burkholderia, which was discussed previously with the betaproteobacteria.)
 Genus Pseudomonas
• In hospitals and other places where pharmaceuGcal agents are
prepared, the ability of pseudomonads to grow on minute
traces of unusual carbon sources, such as soap residues or cap-
liner adhesives found in a soluGon, has been unexpectedly
troublesome.
• Pseudomonads are even capable of growth in some
anGsepGcs, such as quaternary ammonium compounds.
• Their resistance to most an@bio@cs has also been a source of medical
concern.
• This resistance is probably related to the characteris@cs of the cell
wall porins, which control the entrance of molecules through the cell
wall.
• The large genome of pseudomonads also codes for several very
efficient efflux pump systems that eject an@bio@cs from the cell
before they can func@on. Pseudomonads are responsible for about
one in ten nosocomial infec@ons (hospital-acquired infec@ons),
especially among infec@ons in burn units.
• Persons with cysGc fibrosis are also especially prone to
infecGons by Pseudomonas and the closely related
Burkholderia.
 Genus Pseudomonas

• The pseudomonads have a great pracGcal impact in several ways,

including these:
1. Many can degrade an excepGonally wide variety of organic molecules.
• Thus they are very important in the mineraliza)on process (the microbial
breakdown of organic materials to inorganic substances) in nature and in sewage
treatment.
• The fluorescent pseudomonads can use approximately 80 different substances
as their carbon and energy sources.
2. Several species (e.g., P. aeruginosa) are important experimental
subjects.
• Many advances in microbial physiology and biochemistry have come from their
study. For example, the study of P. aeruginosa has significantly advanced our
understanding of how bacteria form biofilms and the role of extracellular
signaling in bacterial communi@es and pathogenesis.
• The genome of P. aeruginosa has an unusually large number of genes for
catabolism, nutrient transport, the efflux of organic molecules, and metabolic
regula@on. This may explain its ability to grow in many environments and resist
an@bio@cs.
 Genus Pseudomonas

3. Some pseudomonads are major animal and plant

pathogens. P. aeruginosa infects people with low
resistance such as cys2c fibrosis pa2ents.
• It also invades burns, and causes urinary tract infec*ons.
P. syringae is an important plant pathogen.
4. Pseudomonads such as P. fluorescens are involved
in the spoilage of refrigerated milk, meat, eggs, and
seafood be- cause 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
demonstraGons of nitrogen fixaGon. 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 mo7le by peritrichous flagella. The cells are ocen pleomorphic, ranging
from rods to coccoid shapes, and form cysts as the culture ages. The genus is aerobic, catalase posi7ve, and
fixes nitrogen non-symbio7cally. 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 conjuncGviGs, an
inflammaGon of the conjuncGva, 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 prepara7ons 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 an7bio7cs. Some
strains are resistant to most available an7bio7cs.
• Not yet widespread in the United States, A. baumanii is an
opportunis7c pathogen primarily found in a hospital sefng.
The an7bio7c resistance of the pathogen, combined with the
weakened health of infected hospital pa7ents, has resulted in
an unusually high mortality rate. A. baumanii is primarily a h;ps://encrypted-tbn0.gstaCc.com/images?q=tbn:ANd9GcRRMFFetaVmc0EV-RKj1jGDqGe99r33SuHJPg&usqp=CAU

respiratory pathogen, but it also infects skin and soc 7ssues

and wounds and occasionally invades the bloodstream.
• It is more environmentally hardy than most gram- nega7ve
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 edi@on of Bergey’s Manual, where both are
placed in the same order, Legionellales.
§ Because the Coxiella share an intracellular lifestyle with the ricke>sial bacteria, they were previously considered
ricke>sial in nature and grouped with them. Legionella bacteria grow readily on suitable ar@ficial 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 isola@on media then available. Aaer
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 rela@vely common in streams, and they colonize such habitats as warm-
water supply lines in hospitals and water in the cooling towers of air condi@oning systems. An ability to survive and
reproduce within aqua@c amoebas oaen makes them difficult to eradicate in water systems.

Genus Coxiella
• Coxiella burneAi (käks-ė-elʹ lä bėr-neʹtē-ē), which causes Q fever, was formerly grouped with the ricke>sia. Like them,
Coxiella bacteria require a mammalian host cell to reproduce.
• Unlike ricke>sias, Coxiella bacteria are not transmi>ed among humans by insect or @ck bites. Although ca>le @cks harbor
the organism, it is most commonly trans- mi>ed by aerosols or contaminated milk. A spore-like body is present in C.
burneAi. This might explain the bacterium’s rela@vely high resistance to the stresses of airborne transmission and heat
treatment.
 Order Vibrionales
§ Members of the order Vibrionales are faculta3vely anaerobic gram-nega3ve
rods. Many are slightly curved. They are found mostly in aqua3c habitats.
§ There are eight genera in the family: Vibrio, Photobacterium, Salinivibrio,
Listonella, Allomonas, Enterovibrio, Catencoccus, and Grimon9a.
§ Several vibrios are important pathogens. Vibrio cholerae causes cholera, and
V. parahaemoly9cus can cause gastroenteri3s in humans following
consump3on of contaminated seafood. V. anguillarum and others are
responsible for fish diseases.

Genus Vibrio
• Members of the genus Vibrio (vibʹrē-ō) are rods that are oFen slightly curved.
• One important pathogen is Vibrio cholerae (kolʹer-ī), the causa3ve agent of
cholera.
• The disease is characterized by a profuse and watery diarrhea. V.
parahaemoly9cus (pa-rä-hē-mō-liʹ3-kus) causes a less serious form of
gastroenteri3s.
• Usually inhabi3ng coastal salt waters, it is transmiMed to humans
mostly by raw or undercooked shellfish.

• Several members of the family are unusual in being bioluminescent. Vibrio fischeri, V. harveyi, and at least two species of Photobacterium are 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 symbio3cally in the luminous organs of fish and squid.
Order Enterobacteriales

• The members of the order Enterobacteriales are faculta7vely anaerobic, gram-nega7ve rods that are, if mo7le,
peritrichously flagellated.
• Morphologically, the rods are straight. This is an important bacterial group, ocen commonly called enterics.
They inhabit the intes7nal tracts of humans and other animals.
• Most enterics are ac7ve 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 gene7c informa7on between cells, which ocen includes an7bio7c
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 intes7nes.
Genus Escherichia

• The bacterial species Escherichia coli is one of the most common

inhabitants of the human intes7nal tract and is probably the most
familiar organism in microbiology.
• Known about the biochemistry and gene7cs of E. coli, and it
con7nues 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 infec7ons, 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 indica7on of fecal
contamina7on. This Photo by Unknown Author is licensed under CC BY-SA
Genus Salmonella

• Almost all members of the genus Salmonella (salʹmön-el-lä) are poten7ally pathogenic.
• Accordingly, there are extensive biochemical and serological tests to clinically isolate and iden7fy
salmonellae.
• Salmonellae are common inhabitants of the intes7nal tracts of many animals, especially poultry and caVle.
Under unsanitary condi7ons, they can contaminate food.

• The nomenclature of the genus Salmonella is unusual. Instead of mul7ple species, members of the genus
Salmonella that are infec7ous to warm-blooded animals can be considered for prac7cal purposes to be a single
species, Salmonella enterica (en-terʹi-kä).
• This species is divided into more than 2400 serovars, that is, serological varieHes. The term serotype is ocen
used to mean the same thing. By way of explana7on of these terms, when salmonellae are injected into
appropriate animals, their flagella, capsules, and cell walls serve as anHgens that cause the animals to form
anHbodies 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 wriVen
as “Salmonella enterica serovar Typhimurium.”
• The conven7on now used by the Centers for Disease Control and Preven7on (CDC) is to spell out the en7re
name at the first men7on and then abbreviate it as, for example, Salmonella Typhimurium. For simplicity,
we will iden7fy 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
nutri7onal advantage in isolated popula7ons with liVle protein nitrogen in their diet.
• The species Klebsiella pneumoniae (nü-mōʹnē-ī) occasionally causes a serious form of pneumonia in humans.

Genus Serra1a
• SerraHa marcescens (ser-räʹ tē-ä mär-sesʹsens) is a bacterial species dis7nguished by its produc7on of red
pigment.
• In hospital situa7ons, the organism can be found on catheters, in saline irriga7on solu7ons, and in other
supposedly sterile solu7ons. Such contamina7on is probably the cause of many urinary and respiratory tract
infec7ons 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 moFlity.
• Periodically, new generaFons of highly moFle swarmer cells develop, and the process is repeated. As a result, a Proteus colony has
the disFncFve appearance of a series of concentric rings.
• This genus of bacteria is implicated in many infecFons of the urinary tract and in wounds.

Genus Yersinia
• Yersinia pes.s (yėr-sinʹē-ä pesʹFs) 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.
Genus Erwinia
• Erwinia (ėr-wiʹnē-ä) species are primarily plant pathogens; some cause plant soc-rot diseases.
• These species produce enzymes that hydrolyze the pec7n 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 infec7ons and hospital-acquired infec7ons.
• 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 domes7c animals. It causes sepsis in
caVle, 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 transmiVed 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
intes7nal 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 cyto- chrome system needed for respira7on, and they
obtain these substances from the heme frac7on, known as the X factor, of blood hemoglobin.
• The culture medium must also supply the ofactor nico7namide adenine dinucleo7de (from either NAD+ or
NADP+), which is known as V factor. Clinical laboratories use tests for the requirement of X and V factors to
iden7fy isolates as Haemophilus species.

• Haemophilus influenzae is responsible for several important diseases. It has been a common cause of meningi7s
in young children and is a frequent cause of earaches.
• Other clinical condi7ons caused by H. influenzae include epiglo77s (a life- threatening condi7on in which
the epiglofs becomes infected and inflamed), sep7c arthri7s in children, bronchi7s, and pneumonia.
Haemophilus ducreyi (dü-krāʹē) is the cause of the sexually transmiVed disease chancroid.
 Class Deltaproteobacteria

• The 𝛅-proteobacteria are disGncGve 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 parFcularly

interesFng genus.
• It aTacks other gram-negaFve bacteria. It
aTaches Fghtly (bdella = leech), and aVer
penetraFng the outer layer of gram-negaFve
bacteria, it reproduces within the periplasm.
• The cell elongates into a Fght 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 intesFnal
tracts of humans and animals.
• Sulfur-reducing and sulfate-reducing bacteria
use organic com- pounds such as lactate,
ethanol, or faTy 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

• Vegeta3ve cells of the myxobacteria (myxo=nasal mucus) move by gliding and leave behind a slime trail.
• Myxococcus xanthus (micks-ō-kokʹkus zanʹthus) and M. fulvus (fulʹvus) are well-studied representa3ves of the genus.
• As they move, their source of nutri3on is the bacteria they encounter, enzyma3cally lyse, and digest.
• Large numbers of these gram-nega3ve microbes eventually aggregate (Figure 11.11).
• Where the moving cells aggregate, they differen3ate and form a macroscopic stalked frui3ng body that contains large numbers of res3ng cells
called myxospores.
• Differen3a3on is usually triggered by low nutrients.
• Under proper condi3ons, usually a change in nutrients, the myxospores germinate and form new vegeta3ve gliding cells.
 Class Epsilonproteobacteria

• The 𝜺-proteobacteria slender gram-nega*ve rods that are helical or curved.

• are the smallest of the five proteobacterial classes.
• They all are slender gram-nega*ve rods, which can be straight, curved, or helical.
• The 𝜺-proteobacteria have one order, Campylobacterales, and three families:
Campylobacteraceae, Helicobacteraceae, and the recently added NauBliaceae.
• Two pathogenic genera, Campylobacter and Helicobacter, are microaerophilic, mo*le,
helical or vibrioid, gram-nega*ve 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 reproduc7ve disease and abor7ons in caVle and sheep. It is associated with a variety of condi7ons in
humans ranging from sep:cemia (pathogens or their toxins in the blood) to enteri:s (inflamma7on of the intes7nal
tract). C. jejuni causes abor7on in sheep and enteri7s diarrhea in humans.

Genus Helicobacter

• Members of the genus Helicobacter are micro-aerophilic curved rods with mul;ple
flagella.
• The species Helicobacter pylori (hēʹlik-ō-bak-tėr pī-lōrʹē) has been iden;fied as the
most common cause of pep;c ulcers in humans and a cause of stomach cancer.
• There are at least 23 species of Helicobacter, all isolated from the stomachs and
upper intes;nes of humans, dogs, cats, and other mammals.
• In developing countries 70 to 90% of the popula;on is infected; developed
countries range from 25 to 50%.
• Most infec;ons are probably acquired during childhood, but the precise
mode of transmission is unclear. The major human pathogen is Helicobacter
pylori, which causes gastri;s and pep;c ulcer disease. H. pylori produces
large quan;;es of urease, and urea hydrolysis appears to be associated with
its virulence.