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UNIT 2

Part 2
PROKARYOTES

Diversity of Bacteria
What does bacterial diversity mean?

Figure 15.1 Major functional traits mapped across major phyla of


Bacteria and Archaea.
The dendrogram shows relationships between microbial phyla as
inferred by analysis of 16S ribosomal RNA gene sequences. Blue
branches are used to denote phyla of Bacteria and red branches
phyla of Archaea. Colored circles indicate phyla that contain at
least one species with a functional trait indicated in the color key.
Metabolic Diversity of Microorganisms
Grupos nutricionales

Fuentes de Energía Fuentes de Carbono

• Fotótrofo: Usa luz • Autótrofo: CO2

• Quimiótrofo: Usa compuestos químicos • Heterótrofo: Compuestos orgánicos

- Orgánicosà Organotrofo • Metilotrofo: Moléculas de C pequeñas


- Inorgánicos à Litotrofo (methanol, metilaminas)
Grupo Fuente de Fuente de
nutricional energía carbono

Fotoautótrofo

Fotoheterótrofo

Quimioautotrofo o
quimiolitótrofo

Quimioheterotrofo o
heterótrofo o
quimiorganotrofo

Mixotrofo
Grupo Fuente de Fuente de Ejemplos
nutricional energía carbono
Cianobacteria, Bacterias
Fotoautótrofo Luz CO2 verdes y púrpuras, algas,
plantas

Fotoheterótrofo Compuestos Algunas bacterias


Luz
orgánicos púrpuras y verdes

Quimioautotrofo o Compuestos Algunas bacterias y


inorgánicos CO2
quimiolitótrofo muchas Archaea
H2, NH3, NO2-, H2S
Quimioheterotrofo o
Compuestos Compuestos La mayoría de las
heterótrofo o
orgánicos orgánicos bacterias, algunas
quimiorganotrofo Archaea y los animales
Compuestos Compuestos
Mixotrofo inorgánicos orgánicos Algunas bacteria
H2 CH3COOH
Relación con el Oxígeno
Þ Aerobios
- obligados: requieren oxígeno (21% o más).
Ej. Bacillus, hongos.
- microaero9licos: requieren niveles menores que el atmosférico (5-10%).
Ej. Azospirillum
- facultativos: no requieren oxígeno, mejor desarrollo con oxígeno.
Ej. levaduras, E. coli
Þ Anaerobios
- aerotolerantes: no son sensibles al oxígeno (crecen en presencia o ausencia de oxígeno).
Ej. Enterococcus faecalis, Sreptococcus spp.
- obligados: no toleran el oxígeno, mueren en su presencia.
Ej. Methanobacterium, Clostridium
Obligate
aerobes

Anaerobes

Facultative
aerobes

Microaerophiles
aerobes

Aerotolerant
anaerobes
Relación con la Temperatura
Diversity of Microorganisms

Phylogenetic diversity: deals with evolutionary


relationships between microorganisms.

Most fundamentally, phylogenetic diversity deals with


the diversity of evolutionary lineages such as phyla,
genera, and species.

Functional diversity is the component of microbial


diversity that deals with diversity in form and function
as it relates to microbial physiology and ecology.
Phylogenetic Diversity of Microorganisms

Over 80 phyla are known

29 major bacterial phyla


(at least one characterized specie in pure culture)
Phylogenetic Diversity of Microorganisms

NOTE: Numbers of cultured and characterized species (green bars) and known 16S rRNA gene
sequences (phylotypes, red bars)
Descending hierarchy of microbial systematics
Bacteria

Bacteria

Proteobacteria

Gammaproteobacteria

Enterobacteriales

Enterobacteriaceae

Escherichia

Escherichia coli
I. Phylum Pseudomonadota
(before Proteobacteria)
Phylum Pseudomonadota (before Proteobacteria)

General characteristics:
Constitute the majority of known bacteria of medical, industrial, and agricultural significance.

All gram-negative bacteria

The largest (1/3) and most metabolically diverse phylum of Bacteria


- wide diversity of energy generating mechanisms chemolithotrophic, chemoorganotrophic,
and phototrophic species.
- diverse in terms of their relationship to oxygen (O2), with anaerobic, microaerophilic, and
facultatively . aerobic species known.

Morphological diverse: exhibit a wide range of cell shapes, including straight and curved rods,
cocci, spirilla, and filamentous, budding, and appendaged forms.
Phylum Pseudomonadota (before Proteobacteria)

Horizontal Gene Flow

Examples:
- Escherichia coli (quimiorganotrofo): model
organism.
- Pseudomonas sp.: capaz de degrader compuestos
tóxicos orgánicos y sintéticos.
- Azotobacter: Nitrogen fixing bacteria
a (alfa) Proteobacteria
General characteristics:
-The second largest class of Pseudomonadota (Proteobacteria)
-One thousand described species
- Most species are obligate aerobes or facultative aerobes.
- Many are oligotrophic à preferring to grow in environments that have low
nutrient concentration.
- A total of 10 orders have been described within the Alphaproteobacteria class
Main orders:
Rhizobiales Domain:
Bacteria
Rickettsiales Kingdom:
Rhodobacterales Bacteria
Rhodospirillales Phylum:
Pseudomonadota
Caulobacterales (Proteobacteria)
Sphingomonadales Calss:
Alphaproteobacteria
Main orders of the
class
Alphaproteobacteria
a (alfa) Proteobacteria: Rhizobiales
The largest and most metabolically diverse order of Alphaproteobacteria contain phototrophs,
chemolithotrophs, symbionts, free-living nitrogen-fixing bacteria a few pathogens of plants and
animals, and diverse chemoorganotrophs

The group gets its name from the rhizobia, a collection of genera that form
root nodules and fix nitrogen in symbiotic association with leguminous
plants.

Domain:
Bacteria
Kingdom:
Bacteria
Phylum:
Pseudomonadota
(Proteobacteria)
Calss:
Alphaproteobacteria
Order:
Rhizobiales
a (alfa) Proteobacteria: Rhizobiales
Rhizobia: typically chemoorganotrophs and obligate aerobes
Each rhizobial genus has a dis[nct range of plant hosts that can be colonized

Nine genera that contain rhizobia:


Bradyrhizobium
Ochrobactrum
Azorhizobium
Devosia
Methylobacterium
Mesorhizobium
Phyllobacterium
Sinorhizobium
and Rhizobium.
a (alfa) Proteobacteria: Rhizobiales
Symbiotic Diazotrophs
Diazotrophs form several symbiotic relationships with plants, animals, and fungi. These relationships are
generally defined by the host providing a hospitable environment, including a source of carbon and energy
and a system for regulating oxygen concentrations, and the microbial symbiont providing in return a supply
of fixed nitrogen to the host.
a (alfa) Proteobacteria: Rhizobiales
Rhizobia can be isolated by crushing nodules and spreading their contents on nutrient-rich solid media;
colonies typically produce copious amounts of exopolysaccharide slime.
a (alfa) Proteobacteria: Rhizobiales
a (alfa) Proteobacteria: Rhizobiales
Agrobacterium tumefaciens (also called Rhizobium radiobacter) is closely related to root nodule
Rhizobium species but is a plant pathogen that causes crown gall disease.

à Insert a plasmid into plant cells, inducing a tumor


Crown gall disease
Overview of the Infection Process
Can be used to transfer DNA directly into the cells of certain plants and fungi.

Encodes genes that mobilize DNA for transfer to the plant.


a (alfa) Proteobacteria: Rhizobiales
The genus Methylobacterium

Is one of the largest in the Rhizobiales.

These species are often called “pink-pigmented facultative methylotrophs” because of


the pink color of their colonies and their good growth on methanol.

Commonly found on the surface of plants and in soils and freshwater systems.
Species of Methylobacterium are readily isolated by pressing the surface of a
plant leaf onto an agar Petri plate containing methanol as the sole source of
carbon.

Figure taken from Virginia Joel et al. (2023)


a (alfa) Proteobacteria: Rhizobiales

These organisms are also commonly encountered in


toilets and baths where their growth on shower
curtains, caulk, and in toilet bowls results in the
formation of pink-pigmented biofilms.

Figures taken from Li N. et al. (2024)


a (alfa) Proteobacteria: Rhizobiales
The Bartonella genus
Intracellular pathogens of humans and other vertebrate animals.

22 species are know, major medical relevance are


• B. henselae
• B. quintana (antes conocida como Rickettsiae quintana)
• B. bacilliformis

B. quintana is the causative agent of trench fever, a disease that


decimated troops in World War I and it´s transmitted to humans
when feces from infected lice are rubbed into abraded skin or the
conjunctiva
a (alfa) Proteobacteria: Rhizobiales
Other species of Bartonella can cause bartonellosis, cat scratch
disease, and a variety of inflammatory diseases.

Disease transmission is mediated by arthropod vectors including


fleas, lice, and sand flies. Species of Bartonella are fas[dious and
difficult to cul[vate, and isola[on is most commonly achieved
using blood agar.
a (alfa) Proteobacteria: Rhizobiales
The genus Pelagibacter
Belongs to the Rhizobiales.

Pelagibacter ubique is an oligotroph and an obligately aerobic


chemoorganotroph that inhabits the photic zone of Earth’s oceans.

This organism can make up 25% of the bacterial cells found at the
ocean’s surface, and its numbers can reach 50% of cells in temperate
waters in the summer; as a consequence, Pelagibacter ubique is likely
the most abundant bacterial species on Earth.

A population of P. ubique cells can double every 29 hours, which is


fairly slow, but they can replicate under low nutrient conditions.
a (alfa) Proteobacteria: Rickettsiales
All obligate intracellular parasites or mutualists of animals.
Species in this order have not yet been cultivated in the absence of host cells.

Those genera that cause disease such as Rickettsia and Ehrlichia are
transmitted by arthropod bites; other genera such as Wolbachia are
obligate parasites or mutualists of insects and other arthropods

Domain:
Bacteria
Kingdom:
Bacteria
Phylum:
Pseudomonadota
(Proteobacteria)
Calss:
Alphaproteobacteria
Order:
Rickettsiales
a (alfa) Proteobacteria: Ricke:siales
Species of the genus Rickettsia are the causative agents of several human diseases (Arthropod-borne).

R. prowazekii à Epidemic typhus


R. typhi à Endemic murine typhus
R. rickettsia à Rocky Mountain Spotted Fever

Metabolically specialized
- Able to oxidize only the amino acids glutamate or
glutamine and unable to oxidize glucose or organic acids.

- Unable to synthesize certain metabolites and must


instead obtain them from host cells
Rickettsias do not survive long outside their hosts, and this may explain why they must be
transmitted from animal to animal by arthropod vectors.
a (alfa) Proteobacteria: Rickettsiales
Rickettsias Growing Within Host Cells

Rickettsia rickettsii in tissue culture.

Once inside the host cell, the bacteria mul[ply primarily


in the cytoplasm and con[nue replica[ng un[l the host
cell is loaded with parasite
a (alfa) Proteobacteria: Rickettsiales
Genus Wolbachia are intracellular parasites of arthropod and insects

Cells of Wolbachia pipientis are passed from an infected female


to her offspring through this egg infection

Affect the reproductive fitness of hosts:


- parthenogenesis (development of unfertilized eggs)
- killing of males
- feminization (the conversion of male insects into
females)
a (alfa) Proteobacteria: Rickettsiales
Genus Wolbachia are intracellular also contain symbionts.

In the wasp Asobara tabida, the development and maturation of females


depend on the presence of Wolbachia.

Without them, the ovaries undergo apoptosis

Huevo de insecto infectado con bacterias simbiontes (Wolbachia sp.)

Credit: Michael Clark & Seth Bordenstein


h-p://discover.mbl.edu/labs.htm
a (alfa) Proteobacteria: Rickettsiales
Genus Ehrlochia à Causal agents of ehrlichiosis a tick-borne
disease (enfermedad transmitida por garrapatas)

figure 31.7 Ehrlichia and Coxiella. (a) Ehrlichia chaffeensis, the causative agent of
human monocytic ehrlichiosis (HME). The electron micrograph shows inclusions in
a human monocyte that contains large numbers of E. chaffeensis cells. The blue
Rash, flulike symptoms, changes in liver function, and a arrows indicate bacteria in each inclusion. The E. chaffeensis cells are about 0.3–0.9
reduction in white blood cell numbers. The symptoms can μm in diameter. Mitochondria are indicated by red arrows.
range from subclinical to fatal.
a (alfa) Proteobacteria: Rickettsiales
TEM of the rickeJsia Coxiella burn/i, cause of Q fever (pneumonia-like infecNon)

C. burnetii cells are transmitted to animals such as sheep, cattle, and goats by insect bites, and from these reservoirs are
transmitted to humans.
figure 31.7 Ehrlichia and Coxiella. (b) Colorized scanning electron micrograph of cells
of Coxiella burnetii, the causative agent of Q fever. The Coxiella cells were grown in
animal cell culture and are shown inside a fractured host cell. A single
C. burnetii cell is about 0.4 μm in diameter.
a (alfa) Proteobacteria: Rhodobacterales and
Rhodospirillales
Metabolically diverse organisms:

-Purple nonsulfur bacteria (Rhodobacter and Rhodospirillum)


-Nitrogen-fixing bacteria (Azospirillum)
-Denitrifiers (Paracoccus), methylotrophs
-Magnetotactic bacteria (Magnetospirillum)

Domain:
Bacteria
Kingdom:
Bacteria
Phylum:
Pseudomonadota
(Proteobacteria)
Calss:
Alphaproteobacteria
Orders:
Rhodobacterales
Rhodospirillales
a (alfa) Proteobacteria: Rhodospirillales
Nitrogen-fixing bacteria:
–Azospirillum spp.
Grow in soil, using nutrients excreted by plants
a (alfa) Proteobacteria: Rhodospirillales
The genus Magnetospirillum

-In a magneNc field, magneNc bacteria demonstrate a dramaNc


directed movement called magnetotaxis. Within these cells are
structures called magnetosomes, which consist of chains of
magneNc parNcles made of magneNte (Fe3O4) or greigite (Fe3S4).

-Magnetosomes are localized within invaginaNons of the cell


membrane that are organized in a linear conformaNon by a protein
scaffold.

-MagneNc bacteria are typically microaerophilic or anaerobic and


are most oVen found near the oxic–anoxic interface in sediments or
straNfied lakes.
a (alfa) Proteobacteria: Caulobacterales
The Caulobacterales are typically oligotrophic (nutrient-
poor) in aquatic environments and strictly aerobic
chemoorganotrophs.

Species typically form prosthecae or stalks

Unequal cell growth

Domain:
Bacteria
Kingdom:
Bacteria
Phylum:
Pseudomonadota
(Proteobacteria)
Calss:
Alphaproteobacteria
Orders:
Caulobacterales
Binary Fission
Growth is the result of cell division and is the
ultimate process in the life of a microbial
cell. In microbiology, growth is defined as an
increase in the number of cells.

When one cell eventually separates to form


two cells, we say that one generation has
occurred, and the time required for this
process is called the generation time.

The generation time of a given bacterial


species is variable and depends on
nutritional and genetic factors, and on
temperature à minutes, hours or days.

Although cell division in most bacteria occurs by binary fission, in a few bacteria other
forms of growth and cell division occur.
a (alfa) Proteobacteria: Caulobacterales
Have prosthecae, are specialized appendages or extensions found in certain microorganisms, can serve various
purposes, including attachment, sensing the environment, cell division, motility or nutrient acquisition.

– Caulobacter –Hyphomicrobium (order rhizobiales)


Stalked bacteria found in lakes (tallos) Budding bacteria (bacterias gemantes) found in lakes
à Order Rhizobiales
Budding bacteria

à Order Caulobacterales
a (alfa) Proteobacteria: Caulobacterales

Domain:
Bacteria
Kingdom:
Bacteria
Phylum:
Pseudomonadota
(Proteobacteria)
Calss:
Alphaproteobacteria Figure 6.24 environmental control of stringent response signaling. b) C. crescentus naturally
Order: inhabits freshwater systems that are limited in nutrients. If cells encounter severe nutrient
limitaMon, the stringent response is induced and ppGpp producMon increases. This increase
Caulobacterales
leads to the cell morphology changing from stalked cells to swarmer cells that can swim to find
more nutrients
The Caulobacter life cycle:

Free-swimming cells alternate with cells that lack flagella and


instead are attached to surfaces by a stalk with a holdfast at
its end.

The role of the swarmer cells is strictly dispersal, as


swarmers cannot divide to form new swarmer cells nor can
they replicate their DNA.

Conversely, the role of the stalked cell is strictly


reproduction. In order to divide, swarmer cells must first
differentiate into stalked cells, and to swim, stalked cells
must first produce swarmers.

Divides into two genetically identical daughter cells that are both
structurally and functionally distinct and express different sets of
genes.
Growth of Caulobacter
Genus
Hyphomicrobium

Domain:
Bacteria
Kingdom:
Bacteria
Phylum:
Pseudomonadota
(Proteobacteria)
Calss:
Alphaproteobacteria
Order:
Rhizobiales
a (alfa) Proteobacteria: Sphingomonadales
Consists of obligately aerobic and nutritionally versatile species

Widespread in aquatic and terrestrial environments and are notable for their ability to metabolize a wide
range of organic compounds including many aromatic compounds that are common environmental
contaminants (e.g., toluene, nonylphenol, dibenzo-p-dioxin, naphthalene, and anthracene, among others)

Domain:
Bacteria
Kingdom:
Bacteria
Phylum:
Pseudomonadota
(Proteobacteria)
Calss:
Alphaproteobacteria
Orders:
Sphingomonadales
a (alfa) Proteobacteria
a (alfa) Proteobacteria
b (beta) Proteobacteria
With about 500 described species, the Betaproteobacteria are the third largest class of
Pseudomonadota (Proteobacteria).
Immense amount of functional diversity
A total of six orders of Betaproteobacteria have many characterized species:

Burkholderiales
Hydrogenophilales
Methylophilales Domain:
Bacteria
Neisseriales Kingdom:
Bacteria
Nitrosomonadales Phylum:
Pseudomonadota
Rhodocyclales (Proteobacteria)
Calss:
Betaproteobacteria
Six major orders
constitute the class
Betaproteobacteria
b (beta) Proteobacteria: Burkholderiales
Contain species with a wide range of metabolic and ecological characteristics

The genus Burkholderia


Metabolic versatility

All species can grow aerobically, some also grow anaerobically with nitrate as the electron acceptor, and
many strains are able to fix N2 à thus some are able to promote plant growth.

Many species are potentially pathogenic for plant or animals.

Nosocomial infections (hospital infection)

Extraordinary nutritional spectrum, able to degrade > 100 different organic molecules, can grow in
disinfectant!
Burkholderia cepacia
complex

Friend or Foe?
b (beta) Proteobacteria: Burkholderiales
One of the best known of the pathogenic species is Burkholderia cepacia.
Soil bacterium but also an opportunisRc pathogen
Can produce both anti-fungal and anti-nematodal compounds, its ability to
colonize plant roots can provide disease protection and promote plant growth

B. cepacia can be pathogenic and it is the major cause of soft rot in onions

Figure 16.8 colonies of Burkholderia. Photograph of colonies of Burkholderia


cepacia on an agar plate.
b (beta) Proteobacteria: Burkholderiales

B. cepacia has also emerged as an opportunistic hospital-acquired infection in


humans, as it is a hardy organism that is difficult to eradicate from the clinical
setting.

Can be a causal agent of secondary lung infections in patients who are


immunocompromised or have pneumonia or cystic fibrosis.

Ability to form biofilms in the lung


Domain:
Natural resistance to many antibiotics
Bacteria
Kingdom:
Bacteria
Phylum: Several reports have cited disinfectants, intravenous solutions and
Pseudomonadota contaminated medical devices as sources of nosocomial Bcc
(Proteobacteria)
Calss:
outbreaks
Betaproteobacteria
Orders:
Burkholderiales
b (beta) Proteobacteria: Burkholderiales
The genus Bordetella

Able to infect the respiratory tracts of mammals

-Bordetella bronchiseptica causes a bronchitis-like cough


in dogs and other domestic animals.

-B. pertussis (more pathogenic than B. bronchiseptica) is


the causative agent of whooping cough (tos ferina), a
serious respiratory disease in humans.
https://doi.org/10.1371/journal.ppat.1007600.g001
https://doi.org/10.1371/journal.ppat.1007600.g001
b (beta) Proteobacteria: Burkholderiales
The genus Sphaerotilus
Common in freshwater environments containing aerobic iron-oxidizers.

They catalyze the oxidation of both iron and manganese.

Sheated microorganisms: form sheaths made of


polysaccharide or protein that encase one or many
Domain:
cells. Sheaths oUen funcRon to bind cells together
Bacteria into long mulRcellular filaments.
Kingdom:
Bacteria
Phylum:
Pseudomonadota
(Proteobacteria)
Calss:
Betaproteobacteria
Orders:
Burkholderiales
b (beta) Proteobacteria: Neisseriales
The order Neisseriales contains at least 29 genera of diverse chemoorganotrophs.
The best-characterized species are in the genera Neisseria and Chromobacterium.

Domain:
Bacteria
Kingdom:
Bacteria
Phylum:
Pseudomonadota
(Proteobacteria)
Calss:
Betaproteobacteria Figure 16.9 Neisseria and Chromobacterium. (a) Transmission electron micrograph of cells of Neisseria
gonorrhoeae showing the typical diplococcus cell arrangements. (b) A large colony of Chromobacterium
Orders: violaceum
Neisseriales
b (beta) Proteobacteria: Neisseriales
The genus Neisseria
Commonly isolated from animals, and some of them are pathogenic.

Neisseria species are always cocci, Chemoheterotrophic

Some Neisseria are free-living saprophytes and reside in the oral


cavity and other moist areas on the animal body and others are
serious human pathogens.
b (beta) Proteobacteria: Neisseriales
The genus Neisseria

-Neisseria gonorrhoeae: the causative agent of the sexually transmitted


disease gonorrhea, adheres specifically to mucosal epithelial cells in the
genitourinary tract, eye, rectum, and throat.

Cells of N. gonorrhoeae are gram-negative diplococci, a morphology not


normally found in microbiota of the urogenital tract. Therefore, a Gram
stain of a urethral, vaginal, or cervical smear revealing such cells, often
surviving inside neutrophils, is diagnostic for gonorrhea. La Mnción de Gram directo del fluido purulento
permite visualizar diplococos intracelulares de
N. gonorrhoeae.
b (beta) Proteobacteria: Neisseriales
Type IV pili key colonization factors for
certain human pathogens, including the
gram-negative N. gonorrhoeae, playing a
key role in attachment to urogenital
epithelia.

N. gonorrhoeae has a cell surface protein


called Opa (opacity associated protein)
that binds specifically to a host protein
found only on the surface of epithelial
cells of these body regions, allowing
adherence of the pathogen to host cells.
b (beta) Proteobacteria: Neisseriales
The genus Neisseria
-Neisseria meningitidis, which can cause a potentially fatal
inflammation of the membranes lining the brain (meningitis).

The bacterium is transmitted to a new host, usually via the


airborne route from an infected individual

Characterized by the sudden onset of a headache accompanied


by vomiting and a stiff neck, and can progress to coma and
death in less than a day.

N. meningitidis bacteremia sometimes leads to fulminant


meningococcemia, a condition characterized by intravascular
coagulation and tissue destruction (gangrene), shock, and
death in over 10% of cases.
b (beta) Proteobacteria: Neisseriales
The genus Chromobacterium
Rod-shaped in morphology.

Facultative aerobe, growing fermentatively on sugars


and aerobically on various carbon sources.

The best-known is Chromobacterium violaceum, a


purple-pigmented organism found in soil and water and
occasionally in pus-forming wounds of humans and
Domain:
other animals. Bacteria
Kingdom:
C. violaceum and other chromobacteria produce the Bacteria
Phylum:
purple pigment violacein a water-insoluble pigment with Pseudomonadota
both antimicrobial and antioxidant properties. (Proteobacteria)
Calss:
Betaproteobacteria
Orders:
Neisseriales
b (beta) Proteobacteria: Rhodocyclales
The order Rhodocyclales contains species with diverse metabolic and ecological characteristics.

The genus Zoogloea

Aerobic chemoorganotrophs that are distinctive for


producing a thick gelatinous capsule that binds cells
together into a complex matrix with branching,
fingerlike projections.

Domain:
Bacteria
Kingdom:
Bacteria
Phylum:
Pseudomonadota
(Proteobacteria)
Calss:
Betaproteobacteria
Orders:
Rhodocyclales
b (beta) Proteobacteria: Rhodocyclales
The order Rhodocyclales contains species with diverse metabolic and ecological characteristics.

The genus Zoogloea


This gelatinous matrix can cause flocculation, the formation
of macroscopic particles that settle out of solution.

Zoogloea ramigera is of particular importance in aerobic


wastewater treatment, where it degrades much of the
organic carbon in the waste stream and promotes
flocculation and settling, crucial steps in water purification.
b (beta) Proteobacteria: Hydrogenophilales, Methylophilales,
and Nitrosomonadales
These three orders contain organisms that have fairly specialized metabolic capabilities including
chemolithotrophs and methylotrophs; most species are obligate aerobes and many are
autotrophic.
Order Hydrogenophilales
Genus Thiobacillus

Commonly found in soils, sulfur springs, marine habitats, and other locales
Domain: where reduced sulfur compounds are available.
Bacteria
Kingdom:
Bacteria chemoorganotrophs
Phylum: chemolithotrophs (oxidize sulfur: H2S ® SO42– )
Pseudomonadota
(Proteobacteria)
Calss: Thiobacillus denitrificans is a classic example, reducing nitrate to dinitrogen gas
Betaproteobacteria
Orders:
Hydrogenophilales
Methylophilales
Nitrosomonadales
b (beta) Proteobacteria: Hydrogenophilales, Methylophilales,
and Nitrosomonadales
These three orders contain organisms that have fairly specialized metabolic capabilities including
chemolithotrophs and methylotrophs; most species are obligate aerobes and many are
autotrophic.
Order Hydrogenophilales
Genus Thiobacillus

Commonly found in soils, sulfur springs, marine habitats, and other locales
Domain: where reduced sulfur compounds are available.
Bacteria
Kingdom:
Bacteria chemoorganotrophs
Phylum: chemolithotrophs (oxidize sulfur: H2S ® SO42– )
Pseudomonadota
(Proteobacteria)
Calss: Thiobacillus denitrificans is a classic example, reducing nitrate to dinitrogen gas
Betaproteobacteria
Orders:
Hydrogenophilales
The b (beta) Proteobacteria
The g (gamma) Proteobacteria
The Gammaproteobacteria are the largest and most diverse class of Proteobacteria, containing
nearly half of all characterized species in the phylum.

The class contains more than 1500 characterized species among its 15 orders
Its species have diverse metabolic and ecological characteristics and include many well-known
human pathogens.

- phototrophic (including the purple sulfur bacteria) Domain:


Bacteria
chemoorganotrophic, or chemolithotrophic Kingdom:
Bacteria
can have either respiratory or fermentative metabolisms. Phylum:
Pseudomonadota
(Proteobacteria)
Calss:
Gammaproteobacteria
12 main orders
consDtute the class
Gammaproteobacteria
The g (gamma) Proteobacteria: Enterobacteriales
Enterobacteriales, one of the largest and best- known orders within the Gammaproteobacteria are
commonly called the enteric bacteria.

Facultatively aerobic, gram-negative, nonsporulating rods (bacilos) that are either nonmotile or motile by
peritrichous flagella.

Enteric bacteria have relatively simple nutritional requirements and ferment


sugars to a variety of end products.
Domain:
Bacteria
Many species pathogenic to humans, other animals, or plants, as well as other Kingdom:
species of industrial importance. Bacteria
Phylum:
Pseudomonadota
Escherichia coli, the best known of all organisms. (Proteobacteria)
Calss:
Gammaproteobacteria
Orders:
Enterobacteriales
The g (gamma) Proteobacteria: Enterobacteriales
The oxidase test and the catalase test are common assays used to characterize bacteria used to
discriminate enteric bacteria from many other Gammaproteobacteria.

Oxidase Test Catalase Test

The catalase test assays for the enzyme


The oxidase test is an assay for the presence
catalase, which detoxifies hydrogen peroxide
of cytochrome c oxidase, an enzyme present
and is commonly found in bacteria able to
in many respiring bacteria
grow in the presence of oxygen

Enteric bacteria are oxidase-negative and catalase-positive.


The g (gamma) Proteobacteria: Enterobacteriales
The oxidase test and the catalase test are common assays used to characterize bacteria used to
discriminate enteric bacteria from many other Gammaproteobacteria.

Oxidase Test Catalase Test

The catalase test assays for the enzyme


The oxidase test is an assay for the presence
catalase, which detoxifies hydrogen peroxide
of cytochrome c oxidase, an enzyme present
and is commonly found in bacteria able to
in many respiring bacteria
grow in the presence of oxygen

Enteric bacteria are oxidase-negative and catalase-positive.


The g (gamma) Proteobacteria: Enterobacteriales
The oxidase test and the catalase test are common assays used to characterize bacteria used to
discriminate enteric bacteria from many other Gammaproteobacteria.

Oxidase Test Catalase Test

Pseudomonas aeruginosa oxidase-negative


Escherichia coli

https://www.youtube.com/watch?v=c5sRWt3DKVw https://www.youtube.com/watch?v=krHqynVe6-g
The g (gamma) Proteobacteria: Enterobacteriales
One major taxonomic characterisRc separaRng the various genera of enteric bacteria is the type and
proporRon of fermentaRon products generated from the fermentaRon of glucose.
Fermentation Patterns in Enteric Bacteria

The mixed-acid fermentation The 2,3-butanediol fermentation


The mixed-acid fermentation
In the mixed-acid fermentaRon, three acids are formed in significant amounts: aceSc, lacSc, and
succinic. Ethanol, CO2, and H2 are also formed, but not butanediol.

Mixed-acid fermentation is observed in


Escherichia
Salmonella
Shigella
Proteus
Citrobacter
Yersinia.
The mixed-acid fermentation
The g (gamma) Proteobacteria: Enterobacteriales
Mixed-Acid Fermenters: The genus Escherichia

-Species of Escherichia are almost universal inhabitants of the intestinal tract of humans and other warm-
blooded animals.

- Play a nutritional role in the intestinal tract by synthesizing vitamins, particularly vitamin K.

- As a facultative aerobe, this organism probably also helps consume O2, thus rendering the large intestine anoxic,
it´s the most abundant facultative anaerobe of the human intestinal microflora.

-Able to grow on a wide variety of carbon and


energy sources such as sugars, amino acids, organic
acids.

- Widely exploited as a cloning host in recombinant


DNA technology.
The g (gamma) Proteobacteria: Enterobacteriales
Mixed-Acid Fermenters: The genus Escherichia

- Some strains are pathogenic and have been implicated in diarrheal diseases
and are major cause of urinary tract infections in women.

Some strains, such as enterohemorrhagic E. coli, an important representative


of which is strain O157:H7, can cause sporadic outbreaks of severe foodborne
disease.

Consump-ion of contaminated foods, such as raw or undercooked ground


beef, unpasteurized milk, or contaminated water. very potent enterotoxin

.
https://doi.org/10.1038/nrmicro818
E. coli produce a variety of colonization factors, many of which are hair-like structures of
various morphologies

Abundant long, straight fimbriae (5–7 nm) of


Fimbria III (long, straight colonization factor antigen)
ETEC contrasRng with thicker, wavy flagella
h-ps://doi.org/10.1038/nrmicro818
Thin (2–3 nm), flexible, wiry CS3 fibrillar structures

P pili showing the thin (∼ 3 nm) fibrillar adhesive tip at


the end of the pilus

https://doi.org/10.1038/nrmicro818
Bundle-forming pilus, aggregates laterally to form large Thin (2–5 nm), coiled, highly aggregative curli
rope-like structures (>10 μm long) of variable width fibres produced by a variety of pathogenic and
non-pathogenic E. coli.
https://doi.org/10.1038/nrmicro818
Pathogenesis of urinary tract infecSon causes by uripathofenic E. coli
Fimbriated strains of Escherichia coli are more frequent causes of urinary tract infections than strains lacking fimbriae

https://doi.org/10.1038/nrmicro818
The g (gamma) Proteobacteria: Enterobacteriales
Mixed-Acid Fermenters: The genus Salmonella

- Closely related to the genus Escherichia

- Salmonella spp. are almost always pathogenic, either to


humans or to other warm-blooded animals (Salmonella is
also found in the intestines of cold-blooded ani- mals, such
as turtles and lizards

In humans typhoid fever and gastroenteritis

Figure 2.18 Fimbriae. Electron micrograph of a dividing cell of Salmonella


enterica (typhi), showing flagella and fimbriae. A single cell is about 0.9 μm wide.
Figure 25.5 Fimbriae. Computer-generated image of a scanning electron micrograph
of cells of Salmonella enterica (typhi) showing the numerous thin fimbriae and the
much thicker peritrichously arranged flagella. A single cell is about 1 μm
in diameter.
“InjecDsome”

Type III systems are commonly used by pathogenic bacteria not only to
secrete toxic proteins outside of the cell but to inject these molecules
directly into eukaryotic host cells.

The entire type III structure is highly complex and


composed of over 100 proteins that facilitate
substrate recognition, coordinated assembly of the
machinery for translocations, and the transport
process itself.

Figure 4.43 Electron micrographs of secretion machinery. Purified


type III injectisomes from Salmonella enterica (typhimurium).
https://www.youtube.com/watch?v=OBf64TEo7gA
Figure 25.10 Virulence factors in Salmonella. Factors important for virulence and the
development of pathogenesis in this gram-negative enteric pathogen are shown.
Genes encoding many of the factors reside on the pathogenicity islands or plasmids
Figure 25.9 Microbial virulence. Differences in microbial virulence
demonstrated by the number of cells of Streptococcus pneumoniae (red
bars) and Salmonella enterica (typhimurium) (green bars) required to
kill mice. Colorized scanning electron micrographs of each bacterium
are shown above their respecave graph.
The g (gamma) Proteobacteria: Enterobacteriales
Mixed-Acid Fermenters: The genus Shigella

- GeneRcally very closely related to Escherichia.


- Genomic analyses strongly suggest that Shigella and Escherichia
have exchanged a significant number of genes by horizontal gene
flow.

- Shigella spp. are typically pathogenic to humans, causing a rather


severe gastroenteriSs called bacillary dysentery.

- Shigella dysenteriae, transmiced by food- and waterborne routes,


which contains endotoxin, invades intesRnal epithelial cells, where it
Figure 5.11 bacterial colonies.
excretes a neurotoxin that causes acute gastrointesRnal distress. (d) Shigella flexneri, grown on
MacConkey agar
The g (gamma) Proteobacteria: Enterobacteriales
Mixed-Acid Fermenters: The genus Proteus

- Typically contains highly motile cells that produce the enzyme urease.

- Frequent cause of urinary tract infections in humans and probably


benefits in this regard from its ready ability to degrade urea by urease.

Proteus mirabilis – swarmer due to


mulRple flagella
Rapid mo5lity of Proteus cells, colonies growing on agar plates o9en exhibit a
characteris5c swarming phenotype

Cells at the edge of the growing colony are more rapidly


motile than those in the center of the colony.

The former move a short distance away from the colony


a mass and then undergo a reduction in motility, settle
down, and divide, forming a new population of motile
cells that again swarm.

As a result, the mature colony appears as a series of


concentric rings, with higher concentrations of cells
alternating with lower concentrations.

Figure 16.13 Swarming in Proteus. (b) Photo of a


swarming colony of Proteus vulgaris. Note the
concentric rings.

Figure 11.9a, b
The 2,3-butanediol fermentation

In the butanediol fermentation, smaller amounts of acids are formed, and butanediol, ethanol, CO2, and
H2 are the main products.

Butanediol fermentation
is characteristic of
Enterobacter
Klebsiella
Erwinia
Serratia
The g (gamma) Proteobacteria: Enterobacteriales
Butanediol fermentation : The genus Serratia

The genus Serratia forms a series of red pyrrole-containing pigments


called prodigiosins.

Prodigiosin is produced in stationary phase as a secondary


metabolite.

Species of SerraUa can be isolated from water and soil as well as from the gut
of various insects and vertebrates and occasionally from the intesRnes of
humans. SerraUa marcescens is also a human pathogen that can cause
infecRons in many body sites. It has been implicated in infecRons caused by
some invasive medical procedures and is an occasional contaminant in
intravenous fluids.
AnRbioRc resistance.
The g (gamma) Proteobacteria: Enterobacteriales
Butanediol fermentation : The genus Serratia

The historian and microbiologist, E. R. L. Gaughran (1969)


uncovered more than 35 historical reports of blood flowing
from Eucharistic bread.
The red pigment offers protection against
excessive UV in sunlight, serves as an
antimicrobial, and has cytotoxic qualities.
The g (gamma) Proteobacteria: Enterobacteriales
Butanediol fermentation : The genus Enterobacter

Enterobacter aerogenes is a common species in water and sewage as well as the intestinal tract of
warm-blooded animals and is an occasional cause of urinary tract infections.

Butanediol fermentation : The genus Klebsiella

Klebsiella pneumoniae, occasionally causes pneumonia in humans, but klebsiellas are most commonly
found in soil and water. Most Klebsiella strains also fix nitrogen, a property not characterisRc of other
enteric bacteria.
The g (gamma) Proteobacteria: Pseudomonadales

Contain exclusively chemoorganotrophs that carry out respiratory metabolisms.

All species can grow as aerobes and are typically oxidase- and catalase-positive, but some are
also capable of anaerobic respiration with nitrate as the electron acceptor.

Most species are able to use a wide diversity of organic compounds as sources of carbon and
energy for growth.
Domain:
Ubiquitous in soil and aquatic systems. Bacteria
Kingdom:
Bacteria
Many species cause diseases of plants and animals, including humans. Phylum:
Pseudomonadota
(Proteobacteria)
Calss:
Gammaproteobacteria
Orders:
Pseudomonadales
The g (gamma) Proteobacteria: Pseudomonadales
The term pseudomonad is often used to describe any gram- negative, polarly flagellated, aerobic rod
that is able to use diverse carbon sources.

Pseudomonads can be found in several different groups of Proteobacteria, but here we consider only
those organisms in the order Pseudomonadales.

Genus:
-Pseudomonas
OpportunisRc
pathogens
Metabolically
diverse
Polar flagella
Figure 16.15 Cell morphology of pseudomonads. Shadow-cast transmission
-Azotobacter and Azomonas: Nitrogen fixing electron micrograph of a Pseudomonas cell. The cell measures about 1 μm in
diameter.
-Moraxella: ConjuncRviRs
The g (gamma) Proteobacteria: Pseudomonadales
The genus Pseudomonas

Several species of Pseudomonas are pathogenic.

- Pseudomonas aeruginosa is frequently associated with infections of the


urinary and respiratory tracts in humans.

- P. aeruginosa is not an obligate pathogen. Instead, the organism is an


opportunist, initiating infections in individuals with weakened immune
systems.
The g (gamma) Proteobacteria: Pseudomonadales
The genus Pseudomonas
P. aeruginosa is one of the most prevalent
pathogens in hospital environments, causing
more than 50% of healthcare-acquired
infections

piocianina (de color azul verdoso), lpioverdina (pigmento fluorescente de


color verde amarillento) y la piorrubina (de color rojo)
The g (gamma) Proteobacteria: Pseudomonadales
The genus Pseudomonas

P. aeruginosa

Colonizes several different surfaces including medical materials


and food industry equipment and form biofilms, leading to
chronic infecRons.

Biofilm production is one of the most important


virulence determinants.

Figure 2. Biofilm formed by Pseudomonas aeruginosa on an orthopedical device. (A)—


Clinical infection after exposure tibial fracture; (B)—Biofilm identification using scanning
electronic microscopy in the orthopedic screw of the same patient; (C,D)—P.
aeruginosa biofilm in the surgical screw under greater magnifications.
https://doi.org/10.3390/pathogens11030300
The g (gamma) Proteobacteria: Pseudomonadales
The genus Pseudomonas
In Pseudomonas aeruginosa, a type VI system is used to
inject a toxin into compeSng bacterial cells.

Type VI systems are widely distributed in gram-negative


bacteria, and like type IV systems they are capable of
delivering a diversity of proteins directly into the
cytoplasm of other cells using a one- step, ATP-requiring
process

Forms a needle-like protein with a pore-forming protein


that contracts all the way through the donor cell’s two
membranes and directly into a host cell once a substrate
molecule is recognized.
The g (gamma) Proteobacteria: Pseudomonadales
The genus Pseudomonas
The g (gamma) Proteobacteria: Vibrionales

Cells are moSle, straight or curved rods


FacultaSve aerobes, with fermentaRve metabolism
Best known genera are Vibrio, Alivibrio, and Photobacterium
Most inhabit aquaRc environments

Domain:
Bacteria
Kingdom:
Bacteria
Phylum:
Pseudomonadota
(Proteobacteria)
Calss:
Gammaproteobacteria
Orders:
Vibrionales
The g (gamma) Proteobacteria: Vibrionales

Vibrio cholerae causes cholera


Some are pathogenic and are found in coastal water
V. parahaemolyticus causes gastroenteritis
The g (gamma) Proteobacteria: Vibrionales
Allivibrio fisheri
Some are capable of light production
Photobacterium phophoreum
(BIOLUMINESCENCE)

Catalyzed by luciferase, an O2-dependent enzyme


Regulation is mediated by population density (quorum sensing)

Although Photobacterium, Aliivibrio, and Vibrio isolates are


facultaRve aerobes, they are bioluminescent only when O2 is
present.

h>ps://www.youtube.com/watch?v=hvd4qPGMj6g
Most bioluminescent bacteria inhabit the marine environment, and some species colonize specialized light
organs of certain marine fishes and squids, producing light that the animal uses for signaling, avoiding
predators, and attracting prey.
Figure 15.47 bioluminescent bacteria and their role as light organ symbionts in the flashlight fish. (a) Two Petri plates of luminous bacteria photographed by
their own light. Note the different colors. Left, Aliivibrio fischeri strain MJ-1, blue light, and (e) (f) right, strain Y-1, green light. (b) Colonies of Photobacterium
phosphoreum photographed by their own light ( Figure 1.2). (c) The flashlight fish Photoblepharon palpebratus; the bright area is the light organ containing
bioluminescent bacteria. (d) Same fish photographed by its own light. (e) Underwater photograph taken at night of P. palpebratus. (f) Electron micrograph of a
thin section through the light-emitting organ of P. palpebratus showing the dense array of bioluminescent bacteria (arrows).
The g (gamma) Proteobacteria
The d (delta) and e (epsilon) Proteobacteria

These classes contain fewer species and less


funcSonal diversity than we have encountered in
the Alpha-, Beta-and Gammaproteobacteria
The d (delta) Proteobacteria
The Delta- proteobacteria are primarily sulfate- and sulfur-reducing bacteria, iron-reducers, and
bacterial predators.

Domain:
Bacteria
Kingdom:
Bacteria
Phylum:
Pseudomonadota
(Proteobacteria)
Calss:
Deltaproteobacteria
The d (delta) Proteobacteria
The Delta- proteobacteria are primarily sulfate- and sulfur-reducing bacteria, dissimilative iron-
reducers, and bacterial predators.

Relevant orders:

Myxococcales and Bdellovibrionales à Contain notable genera of bacterial


predators
Domain:
Bacteria
Kingdom: Desulfuromonadales à Contains diverse species of metal- and sulfur
Bacteria
Phylum: reducing organimsms
Pseudomonadota
(Proteobacteria)
Calss:
Deltaproteobacteria
Orders:
Myxococcales
Bdellovibrionales
The d (delta) Proteobacteria: Bdellovibrionales
The genus Bdellovibrio

Bdellovibrio invade and replicate within the periplasmic space of


their prey cells.
Domain:
Bacteria
Kingdom:
Bacteria
Phylum:
Pseudomonadota
(Proteobacteria)
Calss:
Deltaproteobacteria
Orders:
Figure 15.39 attack on a prey cell by Bdellovibrio. Thin-section electron micrographs of Bdellovibrio attacking
Bdellovibrionales
a cell of Delftia acidovorans. (a) Entry of the predator cell. (b) Bdellovibrio cell inside the host. The Bdellovibrio
cell is enclosed in the bdelloplast and replicates in the periplasmic space. A Bdellovibrio cell measures about
0.3 μm in diameter.
The d (delta) Proteobacteria: Bdellovibrionales
The genus Bdellovibrio

Bdellovibrio are small, highly motile and curved bacteria


that prey on other bacteria, using the cytoplasmic
constituents of their hosts as nutrients.

Obligate aerobe, obtaining its energy from the oxidation of


amino acids and acetate, also assimilates nucleotides, fatty
acids, peptides, and even some intact proteins directly
from its host without first hydrolyzing them.

Widespread in aquatic habitats.


The d (delta) Proteobacteria: Myxococcales
Myxobacteria exhibit the most complex behavioral patterns of all known bacteria
à formation of multicellular structures called fruiting bodies

The fruiting bodies are often strikingly colored and morphologically elaborate can often be seen with a hand
lens on moist pieces of decaying wood or plant material.

Domain:
Bacteria
Kingdom:
Bacteria
Phylum:
Pseudomonadota
(Proteobacteria)
Calss:
Deltaproteobacteria
Orders: Figure 15.41 Fruiting bodies of three species of fruiting myxobacteria. (a) Myxococcus fulvus (125 μm high).
Myxococcales (b) Myxococcus stipitatus (170 μm high). (c) Chondromyces crocatus (560 μm high).
The life cycle of a typical Myxobacterium

Early fru7ng body Mature fruting body -Social Gliding bacteria.


-Cells aggregate to form myxospores
(resting cells)
Specialized cells that are resistant to
drying, UV radiation, and heat, but
the degree of heat resistance is much
less than that of the bacterial
endospore.
Adverse conditions
(nutrient exhaustion)

Figure 15.42 life cycle of Myxococcus xanthus. Aggregation assembles


vegetative cells that then undergo fruiting body formation, within
which some vegetative cells undergo morphogenesis to form resting
cells called myxospores. The myxospores germinate under favorable
nutritional and physical conditions to yield vegetative cells.
The d (delta) Proteobacteria: Myxococcales
Nonflagellated, gram-negative rods that glide across surfaces and
obtain their nutrients primarily by using extracellular enzymes to
lyse other bacteria and use the released nutrients.

Figure 15.43 Myxococcus. (a) Electron micrograph of a thin section of a vegetative cell of Myxococcus xanthus. A cell
measures about 0.75 μm wide. (b) Myxospore of M. xanthus, showing the multilayered outer wall. Myxospores
measure about 2 μm in diameter.
The d (delta) Proteobacteria: Myxococcales
Single coordinated entity in response to environmental cues

Figure 15.45 scanning electron micrographs of fruiRng body formaRon in Chondromyces crocatus. (a) Early stage, showing aggregaaon and
mound formaaon. (b) Iniaal stage of stalk formaaon. Slime formaaon in the head has not yet begun and so the cells that compose the head are
sall visible. (c) Three stages in head formaaon. Note that the diameter of the stalk also increases. (d) Mature fruiang bodies. The enare fruiang
structure is about 600 μm in height (compare with Figure 15.41c).
The d (delta) Proteobacteria: Myxococcales
The d (delta) Proteobacteria: Myxococcales
The e (epsilon) Proteobacteria
Initially defined by only a few pathogenic bacteria.

Environmental studies of marine and terrestrial microbial habitats have shown that a diversity of
Epsilonproteobacteria exist in nature.

Abundant at oxic–anoxic interfaces in sulfur-rich environments, and play major roles in the oxidation of
sulfur compounds in nature.

Representative genus
Campylobacter and Helicobacter

gram- negative, oxidase- and catalase-positive, motile spirilla, and most species are pathogenic to
humans or other animals.

Microaerophilic and must therefore be cultured from clinical specimens at low (3–15%) O2.
The e (epsilon) Proteobacteria
The genus Campylobacter
Acute gastroenteritis that typically results in a bloody diarrhea.

Pathogenesis is due to several factors, including an enterotoxin


that is related to cholera toxin.

Is transmiced to humans via contaminated food


(undercooked poultry or pork, raw shellfish, or occasionally in
fecally contaminated water from surface sources).
Domain:
Bacteria
Kingdom:
Bacteria
Phylum:
Pseudomonadota
(Proteobacteria)
Calss: Figure 32.15 Campylobacter. (a) Colonies of C. jejuni grown on Campylobacter agar, a
Epsilonproteobacteria selective medium. The medium contains several antibiotics to which Campylobacter
Orders: species are naturally resistant. (b) Gram stain and (c) scanning electron micrograph of
Campylobacter cells of a Campylobacter species. Single cells average 0.4 * 2 μm in size.
The e (epsilon) Proteobacteria
The genus Helicobacter

Helicobacter pylori

Pathogen, causes both chronic and acute gastritis, leading to the


formation of peptic ulcers.

Half the world’s population is chronically infected with H. pylori.

Colonizes the surfaces of the gastric mucosa

Chronic inflammation of the gastroduodenum (gastritis) due to


Figure 30.31 Helicobacter pylori. Colorized scanning electron
untreated H. pylori infection may lead to the development of gastric micrograph of cells attached to the mucous lining of the stomach.
cancers. Cells range in length from 3 to 5 μm and are about 0.5 μm in
diameter. Note the flagella.
The e (epsilon) Proteobacteria
The ζ (zeta) Proteobacteria

Is composed of a single known species

The marine iron-oxidizing bacterium Mariprofundus ferrooxydans


Microaerophilic and obligate chemolithotroph
Obtained after the eruption of a volcano

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