CHAPTER 11: THE

THE PROKARYOTIC GROUPS

PROKARYOTES:
 In the second (current) edition of Bergey’s
DOMAINS BACTERIA Manual, the prokaryotes are grouped into
two domains, the Archaea and the
AND ARCHAEA Bacteria.
 Each domain is divided into phyla, each
 In the earlier editions of Bergey’s Manual, phylum into classes, each class into
the main publication that focuses on orders, each order into families, each
classification, bacteria were grouped by family into genera, and finally, each genus
morphology (rod, coccus), staining into species.
reactions, presence of endospores, and  Note that bacteria are also commonly
other obvious features. distinguished by whether they are gram-
 Although this system had its practical uses, negative or gram-positive.
it also had many limitations, somewhat like
grouping bats and birds together on the
basis of their having wings. DOMAIN BACTERIA
 The knowledge of bacteria at the  Actually, relatively few species of bacteria
molecular level has now expanded to such cause disease in humans, animals, plants,
a degree that it is possible to base the or any other organisms.
latest edition of Bergey’s Manual on a
phylogenetic system.
 For example, the genera Rickettsia and
Chlamydia were once grouped together by
their common requirement for intracellular
growth.
 Members of the genus Chlamydia are now
found in a phylum named Chlamydia, but
the rickettsias are now grouped in a distant
phylum, Proteobacteria, in the
Alphaproteobacteria class.
  Bacteria in a symbiotic relationship have
lower metabolic requirements and have the
smallest genomes

GRAM-NEGATIVE The bacterium is extremely small, a little
over 0.3 mm diameter.
BACTERIA  Its small size and minimal genome
probably give it a competitive advantage
PROTEOBACTERIA for survival in low nutrient environments.
 In fact, it seems to be the most abundant
 The proteobacteria, which include most of
living organism in the oceans on the basis
the gram- negative, chemoheterotrophic
of weight.
bacteria, are presumed to have arisen from
 (Part of its name, ubique, is derived from
a common photosynthetic ancestor.
ubiquitous.)
 They are now the largest taxonomic group
 Its sheer numbers must give it an
of bacteria.
important role in the Earth’s carbon cycle.
 However, few are now photosynthetic;
other metabolic and nutritional capacities
Azospirillum
have arisen to replace this characteristic.
 Agricultural microbiologists have been
 The phylogenetic relationship in these
interested in members of the genus
groups is based upon rRNA studies.
Azospirillum, a soil bacterium that grows in
 The name Proteobacteria was taken from
close association with the roots of many
the mythological Greek god Proteus, who
plants, especially tropical grasses.
could assume many shapes.
 It uses nutrients excreted by the plants
 The proteobacteria are separated into five
and in return fixes nitrogen from the
classes designated by Greek letters:
atmosphere.
alphaproteobacteria, betaproteobacteria,
 This form of nitrogen fixation is most
gammaproteobacteria, deltaproteobacteria,
significant in tropical grasses and in
and epsilonproteobacteria
sugarcane, although the organism can be
isolated from the root system of many
The Alphaproteobacteria
temperate- climate plants, such as corn.
 As a group, the alphaproteobacteria
 The prefix azo- is frequently encountered in
includes most of the proteobacteria that are
nitrogen-fixing genera of bacteria.
capable of growth with very low levels of
 It’s derived from a (without) and zo (life), in
nutrients.
reference to the early days of chemistry,
 Some have unusual morphology, including
when oxygen was removed, by a burning
protrusions such as stalks or buds known
candle, from an experimental atmosphere.
as prosthecae.
 Presumably, mammalian life wasn’t
 The alphaproteobacteria also include
possible in this nitrogen-rich atmosphere.
agriculturally important bacteria capable of
 Hence, nitrogen came to be associated
nitrogen fixation in symbiosis with plants,
and several plant and human pathogens with absence of life.
FAMILY Acetobacteraceae
Pelagibacter
 One of the most abundant microorganisms  Genus Acetobacter and Gluconobacter
on Earth, certainly in the ocean are industrially important aerobic
environment, is Pelagibacter ubique organisms that convert ethanol into
 It is a member of a group of marine acetic acid (vinegar).
microbes named SAR 11 because of their  The recently identified Granulibacter is
original discovery in the Sargasso Sea. an emerging pathogen found in patients
 P. ubique is the first member of this group with chronic granulomatous disease.
to be successfully cultivated.
Rickettsia
 Its genome has been sequenced and found
to have only 1354 genes.  In the first edition of Bergey’s Manual, the
genera Rickettsia, Coxiella, and Chlamydia
 This number is very low for a free-living
were grouped closely because they are all
organism, although several mycoplasmas
obligate intracellular parasites—that is,
have even fewer genes.
they reproduce only within a mammalian
cell.
 In the second edition they are now widely  Budding bacteria don’t divide by binary
separated. fission into two nearly identical cells.
 The rickettsias are gram-negative rod-  The budding process resembles the
shaped bacteria, or coccobacilli asexual reproductive processes of many
 One distinguishing feature of most yeasts.
rickettsias is that they are transmitted to  The parent cell retains its identity while the
humans by insect and tick bites, unlike the bud increases in size until it separates as a
Coxiella (discussed later with complete new cell.
gammaproteobacteria).  An example is the genus Hyphomicrobium
 Rickettsia enter their host cell by inducing  These bacteria, like the caulobacteria, are
phagocytosis. found in low-nutrient aquatic environments
 They quickly enter the cytoplasm of the cell and have even been found growing in
and begin reproducing by binary fission laboratory water baths.
 They can usually be cultivated artificially in  Both Caulobacter and Hyphomicrobium
cell culture or chick embryos produce prominent prosthecae.
 The rickettsias are responsible for a
Rhizobium, Bradyrhizobium, and
number of diseases known as the spotted
Agrobacterium
fever group.
 These include epidemic typhus, caused by  The Rhizobium and Bradyrhizobium are two
Rickettsia prowazekii and transmitted by of the more important genera of a group of
lice; endemic murine typhus, caused by R. agriculturally important bacteria that
typhi and transmitted by rat fleas; and specifically infect the roots of leguminous
Rocky Mountain spotted fever, caused by plants, such as beans, peas, or clover.
R. rickettsii and transmitted by ticks  For simplicity these bacteria are known by
 In humans, rickettsial infections damage the common name of rhizobia.
the permeability of blood capillaries, which  The presence of rhizobia in the roots leads
results in a characteristic spotted rash. to formation of nodules in which the rhizobia
and plant form a symbiotic relationship,
Ehrlichia resulting in the fixation of nitrogen from the
 Ehrlichiae are gram-negative, rickettsia-like air for use by the plant
bacteria that live obligately within white  Like rhizobia, the genus Agrobacterium has
blood cells. the ability to invade plants.
 Ehrlichia species are transmitted by ticks to  However, these bacteria do not induce root
humans and cause ehrlichiosis, a nodules or fix nitrogen.
sometimes fatal disease  Of particular interest is Agrobacterium
tumefaciens.
Caulobacter and Hyphomicrobium
 This is a plant pathogen that causes a
 Members of the genus Caulobacter are disease called crown gall; the crown is the
found in low-nutrient aquatic environments, area of the plant where the roots and stem
such as lakes. merge.
 They feature stalks that anchor the  The tumorlike gall is induced when A.
organisms to surfaces tumefaciens inserts a plasmid containing
 This arrangement increases their nutrient bacterial genetic information into the plant’s
uptake because they are exposed to a chromosomal DNA
continuously changing flow of water and  For this reason, microbial geneticists are
because the stalk increases the surface-to- very interested in this organism.
volume ratio of the cell.  Plasmids are the most common vector that
 Also, if the surface to which they anchor is scientists use to carry new genes into a
a living host, these bacteria can use the plant cell because the thick wall of plants is
host’s excretions as nutrients. especially difficult to penetrate
 When the nutrient concentration is
exceptionally low, the size of the stalk Bartonella
increases, evidently to provide an even  The genus Bartonella contains several
greater surface area for nutrient members that are human pathogens.
absorption.
 The best-known is Bartonella henselae, a  There is considerable overlap between the
gram-negative bacillus that causes cat betaproteobacteria and the
scratch disease alphaproteobacteria, for example, among
the nitrifying bacteria discussed earlier.
Brucella  The betaproteobacteria often use nutrient
 Brucella bacteria are small nonmotile substances that diffuse away from areas of
coccobacilli. anaerobic decomposition of organic matter,
 All species of Brucella are obligate parasites such as hydrogen gas, ammonia, and
of mammals and cause the disease methane.
brucellosis  Several important pathogenic bacteria are
 Of medical interest is the ability of Brucella found in this group.
to survive phagocytosis, an important
Acidithiobacillus
element of the body’s defense against
bacteria  Acidithiobacillus species and other sulfur-
oxidizing bacteria are important in the
Nitrobacter and Nitrosomonas sulfur cycle
 Nitrobacter and Nitrosomonas are genera  These chemoautotrophic bacteria are
of nitrifying bacteria that are of great capable of obtaining energy by oxidizing
importance to the environment and to the reduced forms of sulfur, such as
agriculture. hydrogen sulfide (H2S), or elemental sulfur
 They are chemoautotrophs capable of (S0), into sulfates (SO42−).
using inorganic chemicals as energy
Spirillum
sources and carbon dioxide as the only
source of carbon, from which they  The genus Spirillum is found mainly in
synthesize all of their complex chemical freshwater.
makeup.  Spirillum bacteria are motile by conventional
 The energy sources of the genera polar flagella, an important morphological
Nitrobacter and Nitrosomonas (the latter is distinction from the helical spirochetes ,
a member of the betaproteobacteria) are which use axial filaments.
reduced nitrogenous compounds.  The spirilla are relatively large, gram-
 Nitrosomonas species oxidize ammonium negative, aerobic bacteria.
(NH4+) to nitrite (NO2-), which is in turn  Spirillum volutans is often used as a
oxidized by Nitrobacter species to nitrates demonstration slide when microbiology
(NO3-) in the process of nitrification. students are first introduced to the operation
 Nitrate is important to agriculture; it’s a of the microscope
nitrogen form that’s highly mobile in soil
and therefore likely to be encountered and Sphaerotilus
used by plants.  Sheathed bacteria, which include
Wolbachia Sphaerotilus natans are found in
freshwater and in sewage.
 Wolbachia is probably the most common  These gram-negative bacteria with polar
infectious bacterial genus in the world; they flagella form a hollow, filamentous sheath
live only inside the cells of their hosts, in which to live
usually insects (a relationship known as  Sheaths are protective and also aid in
endosymbiosis). nutrient accumulation.
 Therefore, Wolbachia escapes detection by  Sphaerotilus probably contributes to
the usual culture methods. bulking, an important problem in sewage
 Wolbachia interferes with reproduction and treatment
egg development in infected insects.
 Wolbachia-infected male Aedes aegypti Burkholderia
mosquitoes are being released in several  The genus Burkholderia was formerly
places including Brazil, Florida, California, grouped with the genus Pseudomonas,
and southeast Asia to prevent the spread which is now classified under the
of Zika, chikungunya, and dengue viruses. gammaproteobacteria.
The Betaproteobacteria
  Like the pseudomonads, almost all  Members of the order Thiotrichales include
Burkholderia species are motile by a single Thiomargarita namibiensis, which not only
polar flagellum or tuft of flagella. is the largest known bacterium but also
 The best-known species is the aerobic, exhibits several unusual characteristics.
gram-negative rod Burkholderia cepacia  Other members of this order include the
 It has an extraordinary nutritional spectrum nutritionally distinctive genus Beggiatoa
and is capable of degrading more than 100 and Francisella tularensis, the pathogen
different organic molecules. causing tularemia.
 This capability is often a factor in the
Beggiatoa
contamination of equipment and drugs in
hospitals; these bacteria may actually grow  Beggiatoa species are an unusual genus
in disinfectant solutions that grows the only in aquatic sediments at
 This bacterium is also a problem for people the interface between the aerobic and
with the genetic lung disease cystic anaerobic layers.
fibrosis, in whom it metabolizes  Morphologically, the genus resembles
accumulated respiratory secretions. certain filamentous cyanobacteria but it is
 Burkholderia pseudomallei is a resident in not photosynthetic.
moist soils and is the cause of a severe  Its motility is enabled by the production of
disease (melioidosis) endemic in southeast slime, which attaches to the surface on
Asia and northern Australia which movement occurs and provides
lubrication, allowing the organism to glide.
Bordetella
 Nutritionally, Beggiatoa use hydrogen
 Of special importance is the nonmotile, sulfide (H2S) as an energy source and
aerobic, gram-negative rod Bordetella accumulates internal granules of sulfur.
pertussis  The ability of this organism to obtain
 This serious pathogen is the cause of energy from an inorganic compound was
pertussis, or whooping cough an important factor in the discovery of
autotrophic metabolism.
Neisseria
Francisella
 Bacteria of the genus Neisseria are
aerobic, gram-negative cocci that usually  Francisella is a genus of small,
inhabit the mucous membranes of pleomorphic bacteria that grow only on
mammals. complex media enriched with blood or
 Pathogenic species include the tissue extracts.
gonococcus bacterium Neisseria  Francisella tularensis causes the disease
gonorrhoeae, the causative agent of tularemia
gonorrhoea
 and N. meningitidis, the agent of Order Pseudomonadales
meningococcal meningitis  Members of the order Pseudomonadales
are gram-negative aerobic rods or cocci.
Zoogloea
 The most important genus in this group is
 The genus Zoogloea is important in the Pseudomonas.
context of aerobic sewage-treatment  The order also includes Azotobacter,
processes, such as the activated sludge Azomonas, Moraxella, and Acinetobacter.
system.
 As they grow, Zoogloea bacteria form fluffy, Pseudomonas
slimy masses that are essential to the  A very important genus, Pseudomonas
proper operation of such systems. consists of aerobic, gram-negative rods
that are motile by polar flagella, either
The Gammaproteobacteria single or in tufts
 The gammaproteobacteria constitute the  Pseudomonads are very common in soil
largest subgroup of the proteobacteria and and other natural environments.
include a great variety of physiological  Many species of pseudomonads excrete
types. extracellular, water- soluble pigments that
diffuse into their media.
Order Thiotrichales
 One species, Pseudomonas aeruginosa, Pseudomonas and the closely related
produces a soluble, blue-green Burkholderia.
pigmentation.  Although pseudomonads are classified as
 Under certain conditions, particularly in aerobic, some are capable of substituting
weakened hosts, this organism can infect nitrate for oxygen as a terminal electron in
the urinary tract, burns, and wounds, and anaerobic respiration
can cause blood infections, abscesses,  In this way, pseudomonads cause
and meningitis. important losses of valuable nitrogen in
 Other pseudomonads produce soluble fertilizer and soil.
fluorescent pigments that glow when  Nitrate (NO3-) is the form of nitrogen
illuminated by ultraviolet light. fertilizer most easily used by plants.
 One species, P. syringae, is an occasional  Under anaerobic conditions, as in water-
plant pathogen. logged soil, pseudomonads eventually
 (Some species of Pseudomonas have convert this valuable nitrate into nitrogen
been transferred, based upon rRNA gas (N2), which is lost to the atmosphere
studies, to the genus Burkholderia, which  Many pseudomonads can grow at
was discussed previously with the refrigerator temperatures.
betaproteobacteria.)  This characteristic, combined with their
 Although these bacteria are less efficient ability to utilize proteins and lipids, makes
than some other heterotrophic bacteria in them an important contributor to food
utilizing many of the more common spoilage.
nutrients, they compensate for this by
Azotobacter and Azomonas
making use of their genetic capabilities.
 For example, pseudomonads synthesize  Some nitrogen-fixing bacteria, such as
an unusually large number of enzymes and Azotobacter and Azomonas are free-living
can metabolize a wide variety of in soil.
substrates.  These large, ovoid, heavily capsulated
 Therefore, they probably contribute bacteria are frequently used in laboratory
significantly to the decomposition of demonstrations of nitrogen fixation.
uncommon chemicals, such as pesticides,  However, to fix agriculturally significant
that are added to soil. amounts of nitrogen, they would require
 In hospitals and other places where energy sources, such as carbohydrates, that
pharmaceutical agents are prepared, the are in limited supply in soil.
ability of pseudomonads to grow on minute
traces of unusual carbon sources, such as Moraxella
soap residues or capliner adhesives found  Members of the genus Moraxella are strictly
in a solution, has been unexpectedly aerobic coccobacilli—that is, intermediate in
troublesome. shape between cocci and rods.
 Pseudomonads are even capable of  Moraxella lacunata is implicated in
growth in some antiseptics, such as conjunctivitis, an inflammation of the
quaternary ammonium compounds. conjunctiva, the membrane that covers the
 Their resistance to most antibiotics has eye and lines the eyelids
also been a source of medical concern.
 This resistance is probably related to the Acinetobacter
characteristics of the cell wall porins, which  The genus Acinetobacter is aerobic and
control the entrance of molecules through typically forms pairs.
the cell wall  The bacteria occur naturally in soil and
 The large genome of pseudomonads also water. A member of this genus,
codes for several very efficient efflux pump Acinetobacter baumanii is an increasing
systems that eject antibiotics from the cell concern to the medical community because
before they can function. of the rapidity with which it becomes
 Pseudomonads are responsible for about resistant to antibiotics.
one in ten healthcare-associated infections;  Some strains are resistant to most available
especially among infections in burn units. antibiotics.
 People with cystic fibrosis are also  The antibiotic resistance of the pathogen,
especially prone to infections by combined with the weakened health of
 infected patients, has resulted in an  Members of the order Vibrionales are
unusually high mortality rate. facultatively anaerobic gram-negative rods.
 A. baumanii is primarily a respiratory  They are found mostly in aquatic habitats.
pathogen, but it also infects skin, soft  Vibrio species are rods that are often
tissues, and wounds and occasionally slightly curved
invades the bloodstream.  One important pathogen is Vibrio cholerae
 It is more environmentally hardy than most the causative agent of cholera
gram-negative bacteria, and once  The disease is characterized by a profuse
established in a health care setting, it and watery diarrhea.
becomes difficult to eliminate.  V. parahaemolyticus causes a less serious
Legionellales form of gastroenteritis.
 Usually inhabiting coastal salt waters, it’s
 The genera Legionella and Coxiella are transmitted to humans mostly by raw or
closely associated in the second edition of undercooked shellfish.
Bergey’s Manual, where both are placed in
the same order, Legionellales. Enterobacteriales
 Because the Coxiella share an intracellular  The members of the order
lifestyle with the rickettsial bacteria, they Enterobacteriales are facultatively
were previously considered rickettsial in anaerobic, gram-negative rods that are, if
nature and grouped with them. motile, peritrichously flagellated.
 Legionella bacteria grow readily on suitable  Morphologically, the rods are straight.
artificial media  This is an important bacterial group, often
Legionella commonly called enterics.
 This reflects the fact that they inhabit the
 Legionella bacteria were originally isolated intestinal tracts of humans and other
during a search for the cause of an animals.
outbreak of pneumonia now known as  Most enterics are active fermenters of
legionellosis glucose and other carbohydrates.
 The search was difficult because these  Because of the clinical importance of
bacteria didn’t grow on the usual laboratory enterics, there are many techniques to
isolation media then available. isolate and identify them.
 After intensive effort, special media were  Biochemical tests are especially important
developed that enabled researchers to in clinical laboratory work and in food and
isolate and culture the first Legionella. water microbiology.
 Microbes of this genus are now known to  Enterics have fimbriae that help them
be relatively common in streams, and they adhere to surfaces or mucous membranes.
colonize such habitats as warm-water  Specialized sex pili aid in the exchange of
supply lines in hospitals and water in the genetic information between cells, which
cooling towers of air conditioning systems. often includes antibiotic resistance
 An ability to survive and reproduce within  Enterics, like many bacteria, produce
aquatic amebae often makes them difficult proteins called bacteriocins that cause the
to eradicate in water systems. lysis of closely related species of bacteria.
Coxiella  Bacteriocins may help maintain the
ecological balance of various enterics in
 Coxiella burnetii which causes Q fever the intestines.
 Coxiella bacteria require a mammalian host  Important genera of the order include
cell to reproduce. Escherichia, Salmonella, Shigella,
 Coxiella is most commonly transmitted by Klebsiella, Serratia, Proteus, Yersinia,
contaminated milk. Erwinia, Enterobacter, and Cronobacter
 A spore like body is present in C. burnetii
 This might explain the bacterium’s Escherichia
relatively high resistance to the stresses of  The bacterial species Escherichia coli is a
airborne transmission and heat treatment. common inhabitant of the human intestinal
Vibrionales tract and is probably the most familiar
organism in microbiology.
 A great deal is known about the  The convention now used by the Centers
biochemistry and genetics of E. coli, and it for Disease Control and Prevention (CDC)
continues to be an important tool for basic is to spell out the entire name at the first
biological research—many researchers mention and then abbreviate it as, for
consider it almost a laboratory pet. example, Salmonella Typhimurium.
 Its presence in water or food is an  Specific antibodies, which are available
indication of fecal contamination. commercially, can be used to differentiate
 E. coli is not usually pathogenic. Salmonella serovars by a system known as
 However, it can be a cause of urinary tract the Kauffmann-White scheme.
infections, and certain strains produce  This scheme designates an organism by
enterotoxins that cause traveler’s diarrhea numbers and letters that correspond to
and occasionally cause very serious specific antigens on the organism’s
foodborne disease capsule, cell wall, and flagella, which are
 A newly identified emerging pathogenic identified by the letters K, O, and H,
species of E. albertii has been associated respectively.
with sporadic infections in humans, birds,  For example, the antigenic formula for the
and calves. bacterium S. Typhimurium is O1,4,
[5],12:H,i,1,2.*
Salmonella
 Many salmonellae are named only by their
 Almost all members of the genus antigenic formulas
Salmonella are potentially pathogenic.  Serovars can be further differentiated by
 Accordingly, there are extensive special biochemical or physiological
biochemical and serological tests to properties into biovars, or biotypes.
clinically isolate and identify salmonellae.  Typhoid fever, caused by Salmonella Typhi
 Salmonellae are common inhabitants of the is the most severe illness caused by any
intestinal tracts of many animals, especially member of the genus Salmonella
poultry and cattle.  A less severe gastrointestinal disease
 Under unsanitary conditions, they can caused by other S. enterica serovars is
contaminate food. called salmonellosis.
 There are only two species of Salmonella:  Salmonellosis is one of the most common
S. enterica and S. bongori. forms of foodborne illness.
 S. bongori is a resident of “cold-blooded” Shigella
animals—it was originally isolated from a
lizard in the town of Bongor in the African  Species of Shigella are responsible for a
desert nation of Chad—and is rarely found disease called bacillary dysentery, or
in humans. shigellosis
 S. enterica bacteria are infectious to warm-  Unlike salmonellae, they are found only in
blooded animals. humans.
 S. enterica includes more than 2500  Some strains of Shigella can cause life-
serovars, that is, serological varieties. threatening dysentery.
 The term serotype is often used to mean
Klebsiella
the same thing.
 By way of explanation of these terms,  Members of the genus Klebsiella are
when salmonellae are injected into commonly found in soil or water.
appropriate animals, their flagella,  Many isolates are capable of fixing nitrogen
capsules, and cell walls serve as antigens from the atmosphere, which has been
that cause the animals to form antibodies proposed as being a nutritional advantage in
in their blood that are specific for each of isolated populations with little protein
these structures. nitrogen in their diet.
 Thus, serological means are used to  The species Klebsiella pneumoniae
differentiate the microorganisms. occasionally causes a serious form of
 A serovar such as Salmonella pneumonia in humans.
Typhimurium is not a species and should
Serratia
be more properly written as “Salmonella
enterica serovar Typhimurium.”
 Serratia marcescens is a bacterial species  They are widely distributed in humans and
distinguished by its production of red animals, as well as in water, sewage, and
pigment. soil.
 In hospital situations, the organism can be
Cronobacter
found on catheters, in saline irrigation
solutions, and in other supposedly sterile  Cronobacter is a genus of gram- negative,
solutions. rod-shaped bacteria of the family
 Such contamination is probably the cause Enterobacteriaceae.
of many urinary and respiratory tract  This genus was introduced in 2007, and
infections in hospitals. there are now seven named species.
 These bacteria are facultatively anaerobic
Proteus
and generally motile.
 Colonies of Proteus bacteria growing on  The type species is Cronobacter sakazakii
agar exhibit a swarming type of growth. which was previously known as
 Swarmer cells with many flagella move Enterobacter sakazakii.
outward on the edges of the colony and  This organism can cause meningitis and
then revert to normal cells with only a few necrotizing enterocolitis in infants.
flagella and reduced motility.  It is widespread in a range of environments
 Periodically, new generations of highly and foods.
motile swarmer cells develop, and the  Most cases occur in adults, although the
process is repeated. most publicized outbreaks have been
 As a result, a Proteus colony has the associated with contaminated infant
distinctive appearance of a series of formulas.
concentric rings
 This genus of bacteria is implicated in
many infections of the urinary tract and in Pasteurellales
wounds.
 The bacteria in the order Pasteurellales are
Yersinia nonmotile; they are best known as human
and animal pathogens
 Yersinia pestis causes plague, the Black
Death of medieval Europe Pasteurella
 Urban rats in some parts of the world and
ground squirrels in the American  The genus Pasteurella is primarily known
Southwest carry these bacteria. as a pathogen of domestic animals.
 Fleas usually transmit the organisms  It causes sepsis in cattle, fowl cholera in
among animals and to humans, although chickens and other fowl, and pneumonia in
contact with respiratory droplets from several types of animals.
infected animals and people can be  The best-known species is Pasteurella
involved in transmission. multocida which can be transmitted to
humans by dog and cat bites.
Erwinia
Haemophilus
 Erwinia species are primarily plant
pathogens; some cause plant soft-rot  Haemophilus is a very important genus of
diseases. pathogenic bacteria.
 These species produce enzymes that  These organisms inhabit the mucous
hydrolyze the pectin between individual membranes of the upper respiratory tract,
plant cells. mouth, vagina, and intestinal tract.
 This causes the plant cells to separate  The best-known species that affects
from each other, a disease that plant humans is Haemophilus influenzae named
pathologists term soft rot. long ago because of the erroneous belief
that it was responsible for influenza.
Enterobacter  The name Haemophilus is derived from the
 Two Enterobacter species, E. cloacae and bacteria’s requirement for blood in their
culture medium (hemo = blood).
E. aerogenes can cause urinary tract
infections and healthcare-associated  They are unable to synthesize important
infections. parts of the cytochrome system needed for
respiration, and they obtain these
 substances from the heme fraction, known  The product of this reduction is hydrogen
as the X factor, of blood hemoglobin. sulfide (H2S).
 The culture medium must also supply the  (Because the H2S is not assimilated as a
cofactor nicotinamide adenine dinucleotide nutrient, this type of metabolism is termed
(from either NAD+ or NADP+), which is dissimilatory.)
known as V factor.  The activity of these bacteria releases
 Clinical laboratories use tests for the millions of tons of H2S into the atmosphere
requirement of X and V factors to identify every year and plays a key part in the
isolates as Haemophilus species. sulfur cycle
 Haemophilus influenzae is responsible for  Sulfur oxidizing bacteria such as Beggiatoa
several important diseases. are able to use H2S either as part of
 It has been a common cause of meningitis photosynthesis or as an autotrophic energy
in young children and is a frequent cause source.
of earaches.  Desulfovibrio the best-studied sulfur
 Other clinical conditions caused by H. reducing genus, is found in anaerobic
influenzae include epiglottitis (a sediments and in the intestinal tracts of
lifethreatening condition in which the humans and animals.
epiglottis becomes infected and inflamed),  Sulfur-reducing and sulfate-reducing
septic arthritis in children, bronchitis, and bacteria use organic compounds such as
pneumonia. lactate, ethanol, or fatty acids as electron
 Haemophilus ducreyi is the cause of the donors.
sexually transmitted disease chancroid  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
The Deltaproteobacteria
 The deltaproteobacteria are distinctive in
that they include some bacteria that are
Myxococcales
predators on other bacteria.
 Bacteria in this group are also important  In earlier editions of Bergey’s Manual, the
contributors to the sulfur cycle. Myxococcales were classified among the
fruiting and gliding bacteria.
Bdellovibrio
 They illustrate the most complex life cycle
 Bdellovibrio is a particularly interesting of all bacteria, part of which is predatory
genus. upon other bacteria.
 It attacks other gram-negative bacteria.  Vegetative cells of the myxobacteria (myxo
 It attaches tightly (bdella = leech) and after = mucus) move by gliding and leave
penetrating the outer layer of gram- behind a slime trail.
negative bacteria, it reproduces within the  Myxococcus xanthus and M. fulvus are
periplasm. well studied representatives of the genus
 There, the cell elongates into a tight spiral, Myxococcus.
which then fragments almost  As they move, their source of nutrition is
simultaneously into several individual the bacteria they encounter, enzymatically
flagellated cells. lyse, and digest.
 The host cell then lyses, releasing the  Large numbers of these gram- negative
Bdellovibrio cells. microbes eventually aggregate
 Where the moving cells aggregate, they
Desulfovibrionales
differentiate and form a macroscopic
 Members of the order Desulfovibrionales stalked fruiting body that contains large
are sulfur-reducing bacteria. numbers of resting cells called
 They are obligately anaerobic bacteria that myxospores.
use oxidized forms of sulfur, such as  Differentiation is usually triggered by low
sulfates (SO42-) or elemental sulfur (S0) nutrients.
rather than oxygen as electron acceptors.
 Under proper conditions, usually a change  Species that grow in water usually have
in nutrients, the myxospores germinate and gas vacuoles that provide buoyancy,
form new vegetative gliding cells. helping the cell float at a favorable
environment.
The Epsilonproteobacteria
 Cyanobacteria that move about on solid
 The epsilonproteobacteria are slender surfaces use gliding motility.
gram-negative rods that are helical or  Cyanobacteria are morphologically varied.
curved.  They range from unicellular forms that
divide by simple binary fission, to colonial
Campylobacter
forms that divide by multiple fission, to
 Campylobacter bacteria are filamentous forms that reproduce by
microaerophilic vibrios; each cell has one fragmentation of the filaments.
polar flagellum.  The filamentous forms usually exhibit some
 One species, C. fetus causes spontaneous differentiation of cells that are often bound
abortion in domestic animals. together within an envelope or sheath.
 Another species, C. jejuni is a leading  At less than 1 mm, Prochlorococcus is the
cause of outbreaks of foodborne intestinal smallest known photosynthesizer.
disease.  It is one of the most abundant organisms
on Earth, comprising most of the
Helicobacter
photosynthetic population in tropical and
 Helicobacter bacteria are microaerophilic subtropical oceans.
curved rods with multiple flagella.  Prochlorococcus possesses a variety of
 The species Helicobacter pylori has been pigments that enable it to grow as deep as
identified as the most common cause of 200 m.
peptic ulcers in humans and a cause of  Evidence indicates that oxygenic
stomach cancer cyanobacteria played an important part in
the development of life on Earth, which
originally had very little free oxygen that
would support life as we are familiar with it.
 Fossil evidence indicates that when
cyanobacteria first appeared, the
atmosphere contained only about 0.1%
free oxygen.
THE NONPROTEOBACTERIA  When oxygen-producing eukaryotic plants
appeared millions of years later, the
GRAM-NEGATIVE BACTERIA concentration of oxygen was more than
CYANOBACTERIA (THE OXYGENIC 10%.
 The increase presumably was a result of
PHOTOSYNTHETIC BACTERIA)
photosynthetic activity by cyanobacteria.
 The cyanobacteria, named for their  The atmosphere we breathe today contains
characteristic blue-green (cyan) about 20% oxygen.
pigmentation, were once called blue-green  Cyanobacteria occupy environmental
algae. niches similar to those occupied by the
 Although they resemble the eukaryotic eukaryotic algae
algae and often occupy the same  Several cyanobacteria produce toxins that
environmental niches, this is a misnomer sicken humans and other animals
because they are bacteria, algae are not. swimming in water containing large
 However, cyanobacteria do carry out numbers of cyanobacteria
oxygenic photosynthesis, as do the
THE PHYLA CHLOROBI AND
eukaryotic plants and algae
CHLOROFLEXI (THE ANOXYGENIC
 Many of the cyanobacteria are capable of
PHOTOSYNTHETIC BACTERIA)
fixing nitrogen from the atmosphere.
 In most cases, this activity is located in  The phyla Cyanobacteria, Chlorobi, and
specialized cells called heterocysts, which Chloroflexi are gram-negative, but they are
contain enzymes that fix nitrogen gas (N2) not included in the proteobacteria.
into ammonium (NH4+) that can be used
by the growing cell
 Members of the photosynthetic phylum  Members of the phylum Chlamydiae are
Chlorobi (representative genus: grouped with other genetically similar
Chlorobium) are called green sulfur bacteria that do not contain
bacteria. peptidoglycan in their cell walls.
 Members of the phylum Chloroflexi
Chlamydia and Chlamydophila
(representative genus: Chloroflexus) are
called green nonsulfur bacteria.  Chlamydia and Chlamydophila, which we
 Both these varieties of bacteria produce no will call by the common name of the
oxygen during photosynthesis. chlamydias, have a unique developmental
 However, there are photosynthetic gram- cycle that is perhaps their most
negative bacteria that are genetically distinguishing characteristic
included in the proteobacteria.  They are gram-negative coccoid bacteria
 These are the purple sulfur bacteria  Chlamydias are transmitted to humans by
(gammaproteobacteria) and the purple interpersonal contact or by airborne
nonsulfur bacteria (alphaproteobacteria respiratory routes.
and betaproteobacteria).  The chlamydias can be cultivated in
 In these bacterial groups, the term sulfur laboratory animals, in cell cultures, or in the
bacteria indicates that the microbes can yolk sac of embryonated chicken eggs.
use H2S as an electron donor
 If classified as nonsulfur bacteria, the
microbes have at least a limited capability
of phototrophic growth, but without the
production of oxygen.
 Cyanobacteria, as well as eukaryotic plants
and algae, produce oxygen (O2) from
water (H2O) as they carry out
photosynthesis
 The purple sulfur bacteria and green sulfur
bacteria use reduced sulfur compounds,
such as hydrogen sulfide (H2S), instead of
water, and they produce granules of sulfur
(S0) rather than oxygen
 Chromatium is a representative genus.
 At one time, an important question in
biology concerned the source of the
oxygen produced by plant photosynthesis:
was it from CO2 or from H2O?  There are three species of the chlamydias
 Until the introduction of radioisotope that are significant pathogens for humans.
tracers, which traced the oxygen in water  Chlamydia trachomatis is the best-known
and carbon dioxide and settled the pathogen of the group and responsible for
question, comparison of equations 1 and 2 more than one major disease.
was the best evidence that the oxygen  These include trachoma, one of the most
source was H2O. common causes of blindness in humans in
 It is also important to compare these two the less-developed countries
equations to understand how reduced  It is also considered to be the primary
sulfur compounds, such as H2S, can causative agent of both nongonococcal
substitute for H2O in photosynthesis. urethritis, which may be the most common
 Other photoautotrophs, the purple sexually transmitted disease in the United
nonsulfur bacteria and green nonsulfur States, and lymphogranuloma venereum,
bacteria, use organic compounds, such as another sexually transmitted disease
acids and carbohydrates, for the  Two members of the genus Chlamydophila
photosynthetic reduction of carbon dioxide. are well-known pathogens.
 Morphologically, the photosynthetic  Chlamydophila psittaci is the causative
bacteria are very diverse, with spirals, rods, agent of the respiratory disease psittacosis
cocci, and even budding forms. (ornithosis)

PHYLUM Chlamydiae
 Chlamydophila pneumoniae is the cause of  Bacteroides bacteria are gram-negative,
a mild form of pneumonia that is especially are nonmotile, and do not form
prevalent in young adults. endospores.
 Infections caused by Bacteroides often
Planctomycetes
result from puncture wounds or surgery.
 The planctomycetes, a group of gram-  Bacteroides are a frequent cause of
negative, budding bacteria, are said to “blur peritonitis, an inflammation resulting from a
the definition of what bacteria are.” perforated bowel.
 Although their DNA places them among the
Cytophaga
bacteria, they resemble the archaea in the
makeup of their cell walls, and some even  Members of the genus Cytophaga are
have organelles that resemble the nucleus important in the degradation of cellulose
of a eukaryotic cell. and chitin, which are both abundant in soil.
 The members of the genus Planctomyces  Gliding motility places the microbe in close
are aquatic bacteria that produce stalks contact with these substrates, resulting in
resembling Caulobacter and have cell walls very efficient enzymatic action.
similar to those of the archaea, that is,
Fusobacteria
without peptidoglycan.
 One species of planctomycetes, Gemmata  The fusiform bacteria comprise another
obscuriglobus, has a double internal phylum of anaerobes.
membrane around its DNA, resembling a  These bacteria are often pleomorphic but,
eukaryotic nucleus as their name suggests, may be spindle-
shaped (fuso = spindle)
Fusobacterium
 Members of the genus Fusobacterium are
long, slender, gram-negative rods with
pointed rather than blunt ends
 In humans, they are found most often in
the gingival crevice of the gums and may
be responsible for some dental abscesses.
Spirochaetes
 The spirochetes have a coiled morphology,
resembling a metal spring; some are more
tightly coiled than others.
Bacteroidetes  The most distinctive characteristic of this
 The phylum Bacteroidetes includes several phylum, however, is the cells’ method of
genera of aerobic or anaerobic bacteria. motility, which makes use of two or more
axial filaments (or endoflagella) enclosed
 Bacteroidetes are common members of the
in the space between an outer sheath and
human microbiome, especially the
the body of the cell.
gastrointesintal tract.
 One end of each axial filament is attached
 The genus Prevotella is found in the
near one of the cell’s poles
human mouth, and the genus
Elizabethkingia is an emerging cause of  By rotating its axial filament, the cell rotates
healthcare-associated infections. in the opposite direction, like a corkscrew,
which is very efficient in moving the
Bacteroides organism through liquids.
 At the scale of a bacterium, water is as
 Bacteria of the genus Bacteroides live in
viscous as molasses is to a human.
the human intestinal tract in numbers
 However, a bacterium can typically move
approaching 1 billion per gram of feces.
about 100 times its body length in a second
 Some Bacteroides species also reside in
(or about 50 mm/sec); by comparison, a
anaerobic habitats such as the gingival
large, fast fish, such as a tuna, can move
crevice and are also frequently recovered
only about 10 times its body length in this
from deep-tissue infections.
time.
 Many spirochetes are found in the human of the heat-resistant enzyme Taq
oral cavity and are probably among the first polymerase, which is essential to the
microorganisms described by van polymerase chain reaction (PCR).
Leeuwenhoek in the 1600s.  This is the method by which traces of DNA
 An extraordinary location for spirochetes is are amplified and used for identification
on the surfaces of some of the cellulose-
digesting protozoa found in termites, where
they may function as substitutes for flagella.
THE GRAM-POSITIVE
Treponema
BACTERIA
 The gram-positive bacteria can be divided
 The spirochetes include a number of into two groups: those that have a high G +
important pathogenic bacteria. C ratio, and those that have a low G + C
 The best known is the genus Treponema ratio
which includes Treponema pallidum, the  To illustrate the variations in G + C ratio,
cause of syphilis the genus Streptococcus has a low G + C
Borrelia content of 33 to 44%; and the genus
Clostridium has a low content of 21 to 54%.
 Members of the genus Borrelia cause  Included with the gram-positive, low G + C
relapsing fever and Lyme disease serious bacteria are the mycoplasmas, even
diseases that are usually transmitted by though they lack a cell wall and therefore
ticks or lice. do not have a Gram reaction.
Leptospira  Their G + C ratio is 23 to 40%.
 By contrast, filamentous actinomycetes of
 Leptospirosis is a disease usually spread to the genus Streptomyces have a high G +
humans by water contaminated by C content of 69 to 73%.
Leptospira species  Gram-positive bacteria of a more
 The bacteria are excreted in the urine of conventional morphology, such as the
dogs, rats, and swine, so domestic dogs and genera Corynebacterium and
cats are routinely immunized against Mycobacterium, have a G + C content of
leptospirosis. 51 to 63% and 62 to 70%
Deinococcus-Thermus  These bacterial groups are placed into
separate phyla, the Firmicutes (low G + C
 The deinococci include two species of ratios) and Actinobacteria (high G + C
bacteria that have been widely studied ratios).
because of their resistance to extremes in
the environment.
 They stain gram-positive but have a cell wall
that differs slightly in chemical structure from
those of other gram-positives.
 Deinococcus radiodurans is exceptionally
resistant to radiation, even more so than Phylum Firmicutes (Low

endospores.
It can survive exposure to radiation doses as
G + C Gram-Positive
high as 15,000 Grays which is 1500 times
the radiation dosage that would kill a human.
Bacteria)
 The mechanism for this extraordinary  Low G + C gram-positive bacteria are
resistance lies in a unique arrangement of assigned to the phylum Firmicutes.
its DNA that facilitates rapid repair of  This group includes important endospore-
radiation damage. forming bacteria such as the genera
 It is similarly resistant to many mutagenic Clostridium and Bacillus.
chemicals.  Also of extreme importance in medical
 Thermus aquaticus another unique member microbiology are the genera
of this group, is a bacterium that is unusually Staphylococcus, Enterococcus, and
heat stable. Streptococcus.
 It was isolated from a hot spring in
Yellowstone National Park and is the source
 In industrial microbiology, the genus and it didn’t have a membrane-enclosed
Lactobacillus, which produces lactic acid, is nucleus.
well known.  Ribosomal RNA analysis conclusively
 The mycoplasmas, which do not possess a placed Epulopiscium with the prokaryotes.
cell wall, are also found in this phylum.  (The name means “guest at the banquet of
a fish.” It is literally bathed in semi digested
Order Clostridiales
food.)
Clostridium  It most closely resembles gram-positive
bacteria of the genus Clostridium.
 Members of the genus Clostridium are
 Strangely, the species Epulopiscium
obligate anaerobes.
fishelsoni doesn’t reproduce by binary
 The rod-shaped cells contain endospores fission.
that usually distend the cell
 Daughter cells formed within the cell are
 The formation of endospores by bacteria is released through a slit opening in the
important to both medicine and the food parent cell.
industry because of the endospore’s
 This may be related to the evolutionary
resistance to heat and many chemicals.
development of sporulation.
 Diseases associated with clostridia include
 Recently it was discovered that this
tetanus caused by C. tetani.
bacterium does not rely on diffusion to
 botulism caused by C. botulinum and gas distribute nutrients.
gangrene caused by C. perfringens and
 Instead, it makes use of its larger genetic
other clostridia.
capacity—it has 25 times as much DNA as a
 C. perfringens is also the cause of a human cell and as many as 85,000 copies of
common form of foodborne diarrhea. at least one gene—to manufacture proteins
 C. difficile is an inhabitant of the intestinal at internal sites where they are needed.
tract that may cause a serious diarrhea
 This occurs only when antibiotic therapy Order Bacillales
alters the normal intestinal microbiota,  The order Bacillales includes several
allowing overgrowth by toxin-producing C. important genera of gram-positive rods and
difficile. cocci.
Epulopiscium Bacillus
 Biologists have long considered bacteria to  Bacteria of the genus Bacillus are typically
be small by necessity because they lack rods that produce endospores.
the nutrient transport systems used by  They are common in soil, and only a few
higher, eukaryotic organisms and because are pathogenic to humans.
they depend on simple diffusion to obtain
 Several species produce antibiotics.
nutrients.
 Bacillus anthracis causes anthrax, a
 These characteristics would seem to
disease of cattle, sheep, and horses that
critically limit size.
can be transmitted to humans
 So, when a cigar shaped organism living
 It’s often mentioned as a possible agent of
symbiotically in the gut of the Red Sea
biological warfare.
surgeonfish was first observed in 1985, it
 The anthrax bacillus is a nonmotile
was considered to be a protozoan.
facultative anaerobe, often forming chains
 Certainly, its size suggested this: the
in culture.
organism was as large as 80 mm : 600 mm
 The centrally located endospore does not
—over half a millimeter in length—large
distend the walls.
enough to be seen with the unaided eye
 Bacillus thuringiensis is probably the best-
 Compared to the familiar bacterium E. coli,
known microbial insect pathogen
which is about 1 mm : 2 mm, this organism
 It produces intracellular crystals when it
would be about a million times larger in
sporulates.
volume.
 Commercial preparations containing
 Further investigation of the new organism
endospores and crystalline toxin (Bt) of this
showed that certain external structures
bacterium are sold in gardening supply
thought to resemble the cilia of protozoa
shops to be sprayed on plants.
were actually similar to bacterial flagella,
  Bacillus cereus is a common bacterium in  This group includes the principal pathogen
the environment and occasionally is of the streptococci, Streptococcus
identified as a cause of food poisoning, pyogenes also known as the beta-
especially in starchy foods such as rice hemolytic group A streptococcus.
 The three species of the genus Bacillus  Group A represents one of an antigenic
that we have just described are group (A through G) within the hemolytic
dramatically different in important ways, streptococci.
especially their disease-causing properties.  Among the diseases caused by S.
 However, they are so closely related that pyogenes are scarlet fever, pharyngitis
taxonomists consider them to be variants (sore throat), erysipelas, impetigo, and
of a single species. rheumatic fever.
 The most important virulence factor is the
Staphylococcus
M protein on the bacterial surface by which
 Staphylococci typically occur in grapelike the bacteria avoid phagocytosis.
clusters  Another member of the beta-hemolytic
 The most important staphylococcal species streptococci is Streptococcus agalactiae in
is Staphylococcus aureus which is named the beta-hemolytic group B.
for its yellow-pigmented colonies (aureus =  It is the only species with the group B
golden). antigen and is the cause of an important
 Members of this species are facultative disease of the newborn, neonatal sepsis
anaerobes.
Non-beta-hemolytic streptococci
 Some characteristics of the staphylococci
account for their pathogenicity, which takes  Certain streptococci are not beta-
many forms. hemolytic, but when grown on blood agar,
 They grow comparatively well under their colonies are surrounded by a
conditions of high osmotic pressure and distinctive greening.
low moisture, which partially explains why  These are the alpha-hemolytic
they can grow and survive in nasal streptococci.
secretions (many of us carry the bacteria in  The greening represents a partial
our nostrils) and on the skin. destruction of the red blood cells caused
 This also explains how S. aureus can grow mostly by the action of bacteria-produced
in some foods with high osmotic pressure hydrogen peroxide, but it appears only
(such as ham and other cured meats) or in when the bacteria grow in the presence of
low-moisture foods that tend to inhibit the oxygen.
growth of other organisms.  The most important pathogen in this group
 The yellow pigment probably confers some is Streptococcus pneumoniae, the cause of
protection from the antimicrobial effects of pneumococcal pneumonia
sunlight.  Also included among the alpha-hemolytic
 Also, bacterial enzymes digest the fibrin of streptococci are species of streptococci
blood clots, allowing infections to spread called viridans streptococci.
from the sites of injury.  However, not all species form the alpha-
 A few nonpathogenic species of streptococci hemolytic greening (virescent = green), so
are important in the production of dairy this isn’t really a satisfactory group name.
products  Probably the most significant pathogen of
the group is Streptococcus mutans the
primary cause of dental caries

Beta-hemolytic streptococci
 A useful basis for classifying some
streptococci is their colonial appearance Enterococcus
when grown on blood agar.  The enterococci are adapted to areas of
 The beta-hemolytic species produce a the body that are rich in nutrients but low in
hemolysin that forms a clear zone of oxygen, such as the gastrointestinal tract,
hemolysis on blood agar vagina, and oral cavity.
 They are also found in large numbers in
human stool.
 Because they are relatively hardy Lactobacillales but have gradually lost
microbes, they persist as contaminants in a genetic material.
hospital environment, on hands, bedding,  The term degenerative evolution has been
and even as a fecal aerosol. used to describe this process.
 In recent years they have become a  The most significant human pathogen
leading cause of healthcare-associated among the mycoplasmas is M.
infections, especially because of their high pneumoniae which is the cause of a
resistance to most antibiotics. common form of mild pneumonia.
 Two species, Enterococcus faecalis and  Mycoplasmas can be grown on artificial
Enterococcus faecium are responsible for media that provide them with sterols (if
much of the infections of surgical wounds necessary) and other special nutritional or
and the urinary tract. physical requirements.
 In medical settings they frequently enter  Colonies are less than 1 mm in diameter
the bloodstream through invasive and have a characteristic “fried egg”
procedures, such as indwelling catheters appearance when viewed under
magnification
Listeria
 In fact, mycoplasmas grow so well by this
 The pathogenic species of the genus method that they are a frequent
Listeria, Listeria monocytogenes can contamination problem in cell culture
contaminate food, especially dairy laboratories
products.
 Important characteristics of L. Phylum Actinobacteria
monocytogenes are that it survives within
phagocytic cells and is capable of growth (High G + C Gram-
at refrigeration temperatures.
 If it infects a pregnant woman, the Positive Bacteria)
organism poses the threat of stillbirth or
 High G + C gram-positive bacteria are in
serious damage to the fetus.
the phylum Actinobacteria.
Phylum Tenericutes  Many bacteria in this phylum are highly
pleomorphic in their morphology; the
 The Tenericutes phylum includes wall-less genera Corynebacterium and Gardnerella
bacteria called mycoplasmas. for example, and several genera such as
 Once included in the Firmicutes because of Streptomyces grow only as extended,
their low G + C content, mycoplasmas are often-branching filaments.
now in their own phylum, Tenericutes.  Several important pathogenic genera are
 The mycoplasmas are highly pleomorphic found in the Actinobacteria, such as the
because they lack a cell wall and can Mycobacterium species causing
produce filaments that resemble fungi, tuberculosis and leprosy.
hence their name (mykes = fungus, and  The genera Streptomyces, Frankia,
plasma = formed). Actinomyces, and Nocardia are often
 Cells of the genus Mycoplasma are very informally called actinomycetes (from the
small, ranging in size from 0.1 to 0.25 μm, Greek actino = ray) because they have a
with a cell volume that is only about 5% of radiate, or starlike, form of growth by
that of a typical bacillus. reason of their often-branching filaments.
 Because their size and plasticity allowed  Superficially, their morphology resembles
them to pass through filters that retained that of filamentous fungi; however, the
bacteria, they were originally considered to actinomycetes are prokaryotic cells, and
be viruses. their filaments have a diameter much
 Mycoplasmas may represent the smallest smaller than that of the eukaryotic molds.
self-replicating organisms that are capable  Some actinomycetes further resemble
of a free-living existence. molds by their possession of externally
 One species has only 517 genes; the carried asexual spores that are used for
minimum necessary is between 265 and reproduction.
350.  Filamentous bacteria, like filamentous
 Studies of their DNA suggest that they are fungi, are very common inhabitants in soil,
genetically related to the gram-positive
 where a filamentous pattern of growth has  The corynebacteria (coryne = club-shaped)
advantages. tend to be pleomorphic, and their
 The filamentous organism can bridge morphology often varies with the age of the
waterfree gaps between soil particles to cells.
move to a new nutritional site.  The best-known species is
 This morphology also gives the organism a Corynebacterium diphtheriae the causative
much higher surface-to-volume ratio and agent of diphtheria
improves its ability to absorb nutrients in
Propionibacterium
the highly competitive soil environment.
 The name of the genus Propionibacterium
Mycobacterium
is derived from the organism’s ability to
 The mycobacteria are aerobic, non– form propionic acid; some species are
endosporeforming rods. important in the fermentation of Swiss
 The name myco, meaning funguslike, was cheese.
derived from their occasional exhibition of  Propionibacterium acnes are bacteria that
filamentous growth are commonly found on human skin and
 Many of the characteristics of are implicated as the primary bacterial
mycobacteria, such as acid-fast staining, cause of acne.
drug resistance, and pathogenicity, are
Gardnerella
related to their distinctive cell wall, which is
structurally similar to gram-negative  Gardnerella vaginalis is a bacterium that
bacteria causes one of the most common forms of
 However, the outermost lipopolysaccharide vaginitis
layer in mycobacteria is replaced by  There has always been some difficulty in
mycolic acids, which form a waxy, water- assigning a taxonomic position in this
resistant layer. species, which is gram variable and
 This makes the bacteria resistant to exhibits a highly pleomorphic morphology.
stresses such as drying.
Frankia
 Also, few antimicrobial drugs are able to
enter the cell.  The genus Frankia causes nitrogen-fixing
 Nutrients enter the cell through this layer nodules to form in alder tree roots, much
very slowly, which is a factor in the slow as rhizobia cause nodules on the roots of
growth rate of mycobacteria; it sometimes legumes
takes weeks for visible colonies to appear.
Streptomyces
 The mycobacteria include the important
pathogens Mycobacterium tuberculosis  The genus Streptomyces is the best
which causes tuberculosis and M. leprae known of the actinomycetes and is one of
which causes leprosy the bacteria most commonly isolated from
 The mycobacteria are generally separated soil
into two groups:  The reproductive asexual spores of
 (1) the slow growers, such as M. Streptomyces are formed at the ends of
tuberculosis, and aerial filaments.
 (2) the fast, or rapid, growers, which form  If each spore lands on a suitable substrate,
visible colonies on appropriate media it is capable of germinating into a new
within 7 days. colony.
 The slow-growing mycobacteria are more  These organisms are strict aerobes.
likely to be pathogenic to humans.  They often produce extracellular enzymes
 The rapidly growing group also contains a that enable them to utilize proteins,
number of occasional, nontuberculous polysaccharides (such as starch and
human pathogens, which most commonly cellulose), and many other organic
infect wounds. materials found in soil.
 However, these mycobacteria are more  Streptomyces bacteria characteristically
likely to be nonpathogenic soil and water produce a gaseous compound called
microbes. geosmin, which gives fresh soil its typical
musty odor.
Corynebacterium
 Species of Streptomyces are valuable  Most archaea are of conventional
because they produce most of our morphology, that is, rods, cocci, and
commercial antibiotics helices, but some are of very unusual
 This has led to intensive study of the genus morphology
—there are nearly 500 described species.  Some are gram-positive, others gram-
negative; some may divide by binary
Actinomyces fission, others by fragmentation or budding;
 The genus Actinomyces consists of a few lack cell walls.
facultative anaerobes that are found in the  Cultivated members of the archaea
mouth and throat of humans and animals. (singular: archaeon) can be placed into five
 They occasionally form filaments called physiological or nutritional groups.
hyphae that can fragment  Physiologically, archaea are found under
 One species, Actinomyces israelii causes extreme environmental conditions.
actinomycosis, a tissue destroying disease  Extremophiles, as they are known, include
usually affecting the head, neck, or lungs. halophiles, thermophiles, and acidophiles.
 There are no known pathogenic archaea.
Nocardia  Halophiles thrive in salt concentrations of
 The genus Nocardia morphologically more than 25%, such as found in the Great
resembles Actinomyces; however, these Salt Lake and solar evaporating ponds.
bacteria are aerobic.  Examples of these are found in the genus
 To reproduce, they form rudimentary Halobacterium some of which may even
filaments, which fragment into short rods. require such salt concentrations in order to
 The structure of their cell wall resembles grow.
that of the mycobacteria; therefore, they  The optimal growth temperatures of
are often acid-fast. extremely thermophilic archaea is 80°C or
 Nocardia species are common in soil. higher.
 Some species, such as Nocardia  The present record growth temperature
asteroides occasionally cause a chronic, is 121°C, established by archaea
difficult-to-treat pulmonary infection. growing near a hydrothermal vent at
 N. asteroides is also one of the causative 2000 meters deep in the ocean.
agents of mycetoma, a localized  Acidophilic archaea can be found
destructive infection of the feet or hands growing at pH values below zero and
frequently at elevated temperatures, as
DOMAIN ARCHAEA well.
 An example is Sulfolobus whose
 In the late 1970s, a distinctive type of optimal pH is about 2 and optimal
prokaryotic cell was discovered. temperature is more than 70°C.
 Most strikingly, the cell walls of these  Nutritionally, the ocean contains
prokaryotes lacked the peptidoglycan numerous nitrifying archaea that oxidize
common to most bacteria. ammonia for energy.
 It soon became clear that they also shared  Some might also be found in soils.
many rRNA sequences, and the  Methanogens are strictly anaerobic
sequences were different from both those archaea that produce methane as an
of the Domain Bacteria and the Domain end-product by combining hydrogen
Eukarya. (H2) with carbon dioxide (CO2).
 These differences were so significant that  There are no known bacterial
these organisms now constitute a new methanogens.
taxonomic grouping, the Domain Archaea.  These archaea are of considerable
economic importance when they are
used in sewage treatment
 Methanogens are also part of the
microbiota of the human colon, vagina,
and mouth.
DIVERSITY WITHIN THE
ARCHAEA
 although a source of nutritional nitrogen,
MICROBIAL primarily serves as an electron acceptor in
the absence of oxygen.
DIVERSITY  There are reports of bacteria as small as
0.02 to 0.08 mm (nanobacteria) found in
 The Earth provides a seemingly infinite
deep rock formations and even blood
number of environmental niches, and
vessels and kidney stones.
novel life forms have evolved to fill them.
 Most microbiologists have concluded that
 Many of the microbes that exist in these
these are nonliving particles that have
niches cannot be cultivated by
crystallized from minerals.
conventional methods on conventional
 Theoretical considerations have been used
growth media and have remained
to calculate that a cell with a significant
unknown.
metabolism would have to have a diameter
 In recent years, however, isolation and
of at least 0.1 mm.
identification methods have become
 Certain bacteria have extraordinarily small
much more sophisticated, and microbes
genomes.
that fill these niches are being identified
 For example, Carsonella ruddii is a
—many without being cultivated.
bacterium that lives in a symbiotic
 The effects of space travel on bacteria
relationship with its insect host, a sap-
are being studied as the human
eating psyllid (plant louse), and requires
microbiome is now going on spaceflights
less genetic capability than would a free-
DISCOVERIES living microbe.
 It has only 182 genes, which is close to the
ILLUSTRATING THE RANGE 151 genes that is the calculated theoretical
OF DIVERSITY minimum even for a microbe in such a
symbiotic relationship.
 In 1999, another, even larger, giant  C. ruddii is not completely parasitic in its
bacterium was discovered 100 meters relationship with its host insect because it
deep in the sediments of the coastal waters supplies the host with some essential
off Namibia, on the southwestern coast of amino acids.
Africa.  It is therefore probably in the evolutionary
 Named Thiomargarita namibiensis process of becoming an organelle, like the
meaning “sulfur pearl of Namibia,” these mitochondria of mammalian cells
spherical organisms, classified with the  Many bacteria in soil or water, or
gammaproteobacterial, are as large as 750 elsewhere in nature, cannot be cultivated
μm in diameter with the media and conditions normally
 a factor that limits the size of prokaryotic used for bacterial growth.
cells is that nutrients must enter the  Moreover, some bacteria are part of
cytoplasm by simple diffusion. complex food chains and can grow only in
 T. namibiensis minimizes this problem by the presence of other microbes that supply
resembling a fluid-filled balloon, the specific growth requirements.
vacuole in the interior being surrounded by  Recently, researchers have been using the
a relatively thin outer layer of cytoplasm. polymerase chain reaction (PCR) to make
 This cytoplasm is equal in volume to that of millions of copies of genes found at
most other prokaryotes. random in a soil sample.
 Its energy source is essentially hydrogen
sulfide, which is plentiful in the sediments
in which it is normally found, and nitrate,
which it must extract intermittently from
nitrate-rich seawaters when storms stir the
loose sediment.
 The cell’s interior vacuole, which makes up
about 98% of the bacterium’s volume,
serves as a storage space to hold the
nitrate between recharging of its supply.
 The cell’s energy is derived from the
oxidation of hydrogen sulfide; the nitrate,