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EXTREMOPHILES
NATURE’S ULTIMATE SURVIVORS
EXTREMOPHILES

I.      What are they?
II.     Types of Extremophiles
III.    Extreme Prokaryotes
IV.     Extreme Eukaryotes
V.      Extreme Viruses
VI.     Evolution of Extremophiles
VII.    Biotechnological Uses
VIII.   Industrial Uses
IX.     Extraterrestrial Extremophiles?
What are Extremophiles?


Extremophiles are microorganisms— whether
  viruses, prokaryotes, or eukaryotes— that
survive under harsh environmental conditions
  that can include atypical temperature, pH,
 salinity, pressure, nutrient, oxic, water, and
                radiation levels
Types of Extremophiles
Types of Extremophiles
Types of Extremophiles
Other types include:
 Barophiles -survive under high pressure levels, especially
  in deep sea vents
 Osmophiles –survive in high sugar environments
 Xerophiles -survive in hot deserts where water is scarce
 Anaerobes -survive in habitats lacking oxygen
 Microaerophiles -survive under low-oxygen conditions only
 Endoliths –dwell in rocks and caves
 Toxitolerants -organisms able to withstand high levels of
  damaging agents. For example, living in water saturated
  with benzene, or in the water-core of a nuclear reactor
Surviving the Extremes
EXTREME PROKARYOTES
   Hyperthermophiles
              -Members of
              domains Bacteria
              and Archaea
              -Held by many
              scientists to have
              been the earliest
              organisms
              -Early earth was
              excessively hot,
              so these
              organisms would
              have been able to
Morphology of Hyperthermophiles
-Heat stable proteins that have more
   hydrophobic interiors, which prevents
   unfolding or denaturation at higher
   temperatures
-Have chaperonin proteins that maintain
   folding
-Monolayer membranes of dibiphytanyl
   tetraethers, consisting of saturated fatty
   acids which confer rigidity, preventing
   them from being degraded in high
   temperatures
-Have a variety of DNA-preserving substances
   that reduce mutations and damage to
   nucleic acids, such as reverse DNA gyrase The red on these rocks
   and Sac7d                                 is produced by
-They can live without sunlight or organic      Sulfolobus solfataricus,
   carbon as food, and instead survive on       near Naples, Italy
   sulfur, hydrogen, and other materials that
   other organisms cannot metabolize
N acetyltalosaminuuronic acid
a | Schematic representation of a cross-section of the cell envelope of Sulfolobus solfataricus showing the cytoplasmic membrane, with
membrane-spanning tetraether lipids and an S-layer composed of two proteins — a surface-covering protein (red oval) and a membrane-
anchoring protein (yellow oblong). b | Schematic representation of a cell envelope of an archaeon that stains positive with the Gram stain
and that contains a pseudomurein layer in addition to the S-layer. The cytoplasmic membrane is composed of diether lipids.
Some Hyperthermophiles

                              Frequent habitats
                              include volcanic vents
                              and hot springs, as in
                              the image to the left




Pyrococcus abyssi   1µm   Thermus aquaticus   1µm
Deep Sea Extremophiles
                             The deep-sea floor and hydrothermal
                                vents involve the following conditions:
                             low temperatures (2-3º C) – where only
                                psychrophiles are present
                             low nutrient levels – where only
                                oligotrophs present
                             high pressures – which increase at the
                                rate of 1 atm for every 10 meters in
                                depth (as we have learned, increased
                                pressure leads to decreased enzyme-
                                substrate binding)
                                     barotolerant microorganisms live
                                at 1000-4000 meters
A black smoker, a submarine          barophilic microorganisms live at
 hot spring, which can reach    depths greater than 4000 meters
  518- 716°F (270-380°C)
Extremophiles of
                   Hydrothermal Vents
                                                       Natural
                                                       springs which
                                                       vent warm or
                                                       hot water on
                                                       the sea floor
                                                       near mid-
                                                       ocean ridges.
                                                        Associated
                                                       with the
                                                       spreading of
    0.2µm                        1µm                   the earth’s
A cross-section of a bacterium     A bacterial         crust. High
isolated from a vent. Often        community from a    temperatures
such bacteria are filled with      deep-sea            and pressures
viral particles which are          hydrothermal vent
abundant in hydrothermal           near the Azores      
vents
Psychrophiles




                                     Some microorganisms
                                     thrive in temperatures
                                          well below the
                                         freezing point of
                                        water, such as in
     Some researchers believe that          Antarctica
psychrophiles live in conditions mirroring
         those found on Mars
Psychrophiles possess:

  -proteins rich in α-helices and polar groups which allow for
  greater flexibility


  -“antifreeze proteins” that maintain liquid intracellular conditions
  by lowering freezing points of other biomolecules

  -membranes that are more fluid, containing unsaturated cis-fatty
  acids which help to prevent freezing

  -active transport at lower temperatures
Halophiles




-Divided into mild (1-6%NaCl), moderate (6-15%NaCl), and extreme
(15-30%NaCl)

-Halophiles are mostly obligate aerobic archaea

How do halophiles survive high salt concentrations?
-by interacting more strongly with water such as using more
negatively charged amino acids in key structures
-by making many small proteins inside the cell, and these, then,
compete for the water
-and by accumulating high levels of salt in the cell in order to
outweigh the salt outside
Barophiles
                         -Survive under levels
                         of pressure that are
                         otherwise lethal to
                         other organisms
                         -Usually found deep in
                         the earth, in the deep
                         sea, hydrothermal
                         vents, etc
1µm                      -scientists believe that
A sample of barophilic   barophiles may be
bacteria from the        able to survive on the
earth’s interior         Moon and other places
                         in space
Xerophiles
 Extremophiles which live in water-
  scarce habitats, such as deserts


 Produce desert varnish as seen
     in the image to the left


 Desert varnish is a thin coating of
 Mn, Fe, and clay on the surface of
  desert rocks, formed by colonies
    of bacteria living on the rock
   surface for thousands of years
SOME COMMON GENERA OF PROKARYOTE EXTREMOPHILES




2um              1.8um             1um

  Thermotoga             Aquifex    Halobacterium




0.6um            0.9um             0.9um
Methanosarcina     Thermoplasma     Thermococcus




1.3um            0.6um
                                     0.7um
Thermoproteus        Pyrodictium         Ignicoccus
Deinococcus radiodurans
                 The Radiation Resistor
                                -Possesses extreme resistance to
                                up to 4 million rad of radiation,
                                genotoxic chemicals (those that
                                harm DNA), oxidative damage from
                                peroxides/superoxides, high levels
                                of ionizing and ultraviolet
                                radiation, and dehydration
                                -It has from four to ten DNA
                                molecules compared to only one
                                for most other bacteria
  0.8µm

-Contains many DNA repair enzymes, such as RecA, which
matches the shattered pieces of DNA and splices them back
together. During these repairs, cell-building activities are shut off
and the broken DNA pieces are kept in place
Chroococcidiopsis
   The Cosmopolitan Extremophile




         1.5µm

-A cyanobacteria which can survive in a variety of harsh
environments, such as hot springs, hypersaline habitats, hot,
arid deserts throughout the world, and in the frigid Ross
Desert in Antarctica
-Possesses a variety of enzymes which assist in such
adaptation
Other Prokaryotic Extremophiles




 1µm                           1µm
Gallionella ferrugineaand      Anaerobic bacteria
(iron bacteria), from a cave



 Current efforts in microbial taxonomy are isolating more and
 more previously undiscovered extremophile species, in places
 where life was least expected
EXTREME EUKARYOTES
  THERMOPHILES/ACIDOPHILES




                             2µm
EXTREME EUKARYOTES
                          PSYCHROPHILES




       2µm

Snow Algae (Chlamydomonas nivalis) A bloom of Chloromonas
                                   rubroleosa in Antarctica
     These algae have successfully adapted to their harsh
     environment through the development of a number of
  adaptive features which include pigments to protect against
   high light, polyols (sugar alcohols, e.g. glycerine), sugars
  and lipids (oils), mucilage sheaths, motile stages and spore
EXTREME EUKARYOTES
                         ENDOLITHS

                                    Quartzite from Johnson
                                    Canyon, California.
                                    Sample shows green
                                    bands of endolithic
                                    algae. Rock is 9.5 cm
                                    wide




-Endoliths (also called hypoliths) are usually
algae, but can also be prokaryotic cyanobacteria,
that exist within rocks and caves
-Often are exposed to anoxic (no oxygen) and
anhydric (no water) environments
EXTREME EUKARYOTES
                      PARASITES
 -Members of the Phylum Protozoa, which are
   regarded as the earliest eukaryotes to evolve, are
   mostly parasites
 -Parasitism is often a stressful relationship on both
   host and parasite, so they are considered
   extremophiles




                                20µm
  15µm

Trypanosoma gambiense,         Balantidium coli, causes
causes African sleeping        dysentery-like symptoms
sickness
EXTREME VIRUSES
                             Viruses are currently being
                            isolated from habitats where
                             temperatures exceed 200°F


                                Instead of the usual
                            icosahedral or rod-shaped
                            capsids that known viruses
                            possess, researchers have
   40nm
                             found viruses with novel
Virus-like particles         propeller-like structures
isolated from the extreme
environment of
Yellowstone National Park   These extreme viruses often
hot springs                   live in hyperthermophile
                                 prokaryotes such as
                                      Sulfolobus
CLASSIFICATION OF EXTREMOPHILES

   Phylogenetic Relationships




 Extremophiles are present among Bacteria, form
the majority of Archaea, and also a few among the
                     Eukarya
PHYLOGENETIC RELATIONSHIPS
-Members of Domain Bacteria (such as Aquifex and
  Thermotoga) that are closer to the root of the
  “tree of life” tend to be hyperthermophilic
  extremophiles
-The Domain Archaea contain a multitude of
  extremophilic species:
  Phylum Euryarchaeota-consists of methanogens
  and extreme halophiles
  Phylum Crenarchaeota-consists of
  thermoacidophiles, which are extremophiles that
  live in hot, sulfur-rich, and acidic solfatara
  springs
  Phylum Korarchaeota-new phylum of yet
  uncultured archaea near the root of the Archaea
  branch, all are hyperthermophiles
-Most extremophilic members of the Domain
  Eukarya are red and green algae
Chronology of Life
The First Organisms?
Early Earth was largely inhospitable: high CO 2/H2S/H2
   etc, low oxygen, and high temperatures
Lifeforms that could evolve in such an environment
   needed to adapt to these extreme conditions
H2 was present in abundance in the early
   atmosphere. Many hyperthermophiles and
   archaea are H2 oxidizers
Thus, it is widely held that extremophiles represent
   the earliest forms of life with non-extreme forms
   evolving after cyanobacteria had accumulated
   enough O2 in the atmosphere
Results of rRNA and other molecular techniques
   have placed hyperthermophilic bacteria and
   archaea at the roots of the phylogenetic tree of
   life
Evolutionary Theories


 Consortia- symbiotic relationships between microorganisms,
  allows more than one species to exist in extreme habitats
  because one species provides nutrients to the others and vice
  versa
 Genetic drift appears to have played a major role in how
  extremophiles evolved, with allele frequencies randomly
  changing in a microbial population. So alleles that conferred
  adaptation to harsh habitats increased in the population, giving
  rise to extremophile populations
 Geographic isolation may also be a significant factor in
  extremophile evolution. Microorganisms that became isolated in
  more extreme areas may have evolved biochemical and
  morphological characteristics which enhanced survival as
  opposed to their relatives in more temperate areas. This
  involves genetic drift as well
Slower Evolution
-Extremophiles, especially hyperthermophiles,
  possess slow “evolutionary clocks”
-That is, they have not evolved much from their
  ancestors as compared to other organisms
-Hence, hyperthermophiles today are similar to
  hyperthermophiles of over 3 billion years ago
-Slower evolution may be the direct result of living
  in extreme habitats and little competition
-By contrast, other extremophiles, such as extreme
  halophiles and psychrophiles, appear to have
  undergone faster modes of evolution since they
  live in more specialized habitats that are not
  representative of early earth conditions
Mat Consortia
   Mat Consortia


                                                A mat
                                                consortia in
                                                Yellowstone




-Microbial mats consist of an upper layer of photosynthetic
bacteria, with a lower layer of nonphotosynthetic bacteria
-These consortia may explain some of the evolution that has
taken place: extremophiles may have relied on other
extremophiles and non-extremophiles for nutrients and shelter
-Hence, evolution could have been cooperative
USES OF EXTREMOPHILES
HYPERTHERMOPHILES (SOURCE)            USES
DNA polymerases                       DNA amplification by PCR
Alkaline phosphatase                  Diagnostics
Proteases and lipases                 Dairy products
Lipases, pullulanases and proteases   Detergents
Proteases                             Baking and brewing and amino
                                      acid production from keratin
Amylases, α-glucosidase, pullulanase and xylose/glucose isomerases
                                       Baking and brewing and amino
                                       acid production from keratin
Alcohol dehydrogenase                 Chemical synthesis
Xylanases                             Paper bleaching
Lenthionin                            Pharmaceutical
S-layer proteins and lipids           Molecular sieves
Oil degrading microorganisms          Surfactants for oil recovery
Sulfur oxidizing microorganisms       Bioleaching, coal & waste gas
                                      desulfurization
Hyperthermophilic consortia           Waste treatment and methane
                                      production
USES OF EXTREMOPHILES
PSYCHROPHILES (SOURCE)           USES
Alkaline phosphatase             Molecular biology
Proteases, lipases, cellulases and amylases
                                 Detergents
Lipases and proteases            Cheese manufacture and dairy
                                 production
Proteases                        Contact-lens cleaning solutions,
                                 meat tenderizing
Polyunsaturated fatty acids      Food additives, dietary
                                 supplements
Various enzymes                  Modifying flavors
b-galactosidase                  Lactose hydrolysis in milk
                                 products
Ice nucleating proteins       Artificial snow, ice cream, other
                              freezing applications in the food
                              industry
Ice minus microorganisms      Frost protectants for
                              sensitive plants
Various enzymes (e.g. dehydrogenases)
                              Biotransformations
Various enzymes (e.g. oxidases)Bioremediation, environmental
                               biosensors
Methanogens                    Methane production
USES OF EXTREMOPHILES
HALOPHILES (SOURCE)                 USES
Bacteriorhodopsin                   Optical switches and photocurrent generators in
                                    bioelectronics
Polyhydroxyalkanoates               Medical plastics
Rheological polymers                Oil recovery
Eukaryotic homologues (e.g. myc oncogene product)
                                    Cancer detection, screening anti-tumor drugs
Lipids                              Liposomes for drug delivery and cosmetic
                                    packaging
Lipids                              Heating oil
Compatible solutes                  Protein and cell protectants in variety of
                                    industrial uses, e.g. freezing, heating
Various enzymes, e.g. nucleases, amylases, proteases
                                    Various industrial uses, e.g. flavoring agents
g-linoleic acid, b-carotene and cell extracts, e.g. Spirulina and Dunaliella
                                    Health foods, dietary supplements, food coloring
                                    and feedstock
Microorganisms                      Fermenting fish sauces and modifying food
                                    textures and flavors
Microorganisms                      Waste transformation and degradation, e.g.
                                    hypersaline waste brines contaminated with a
                                    wide range of organics
Membranes                           Surfactants for pharmaceuticals
USES OF EXTREMOPHILES

ALKALIPHILES (SOURCE)                    USES
Proteases, cellulases, xylanases, lipases and pullulanases
                                         Detergents
Proteases                                Gelatin removal on X-ray
                                         film
Elastases, keritinases                   Hide dehairing
Cyclodextrins                            Foodstuffs, chemicals and
                                         pharmaceuticals
Xylanases and proteases                  Pulp bleaching
Pectinases                               Fine papers, waste
                                         treatment and degumming
Alkaliphilic halophiles                  Oil recovery
Various microorganisms                   Antibiotics



ACIDOPHILES (SOURCE)                   USES
Sulfur oxidizing microorganisms        Recovery of metals and
                               desulfurication of coal
Microorganisms                         Organic acids and solvents
Taq Polymerase
     Isolated from the
     hyperthermophile
     Thermus aquaticus
     Much more heat stable
     Used as the DNA
     polymerase in the very
     useful Polymerase
     Chain Reaction (PCR)
     technique which
     amplifies DNA samples
Alcohol Dehydrogenase
   -Alcohol dehydrogenase (ADH), is
   derived from a member of the archaea
   called Sulfolobus solfataricus
   -It works under some of nature's
   harshest volcanic conditions: It can
   survive to 88°C (190ºF) - nearly boiling
   - and corrosive acid conditions
   (pH=3.5) approaching the sulfuric acid
   found in a car battery (pH=2)
   -ADH catalyzes the conversion of
   alcohols and has considerable
   potential for biotechnology
   applications due to its stability under
   these extreme conditions
Bacteriorhodopsin
        -Bacteriorhodopsin is a
        trans-membrane protein
        found in the cellular
        membrane of
        Halobacterium
        salinarium, which
        functions as a light-
        driven proton pump


        -Can be used for
        electrical generation
Bioremediation
-    Bioremediation is the branch of biotechnology
     that uses biological processes to overcome
     environmental problems

-    Bioremediation is often used to degrade
     xenobiotics introduced into the environment
     through human error or negligence

    - Part of the cleanup effort after the 1989
      Exxon Valdez oil spill included
      microorganisms induced to grow via nitrogen
      enrichment of the contaminated soil
Bioremediation
Psychrophiles as Bioremediators
-   Bioremediation applications with cold-
    adapted enzymes are being considered for
    the degradation of diesel oil and
    polychlorinated biphenyls (PCBs)

- Health effects that have been associated with
  exposure to PCBs include acne-like skin
  conditions in adults and neurobehavioral and
  immunological changes in children. PCBs are
  known to cause cancer in animals
An End to Pollution?


New and innovative methods are being
developed that utilize extremophiles for the
elimination of pollution resulting from oil
slicks, toxic chemical spills, derelict mines,
etc
Life in Outer Space?
-Scientists have decided on 3 requirements for life:
        water
        energy
        carbon
-Astrobiology: field of biology dealing with the existence of life
   beyond earth
-Astrobiologists are currently looking for life on Mars,
   Jupiter’s moon Europa, and Saturn’s moon Titan
-Such life is believed to consist of extremophiles that can
   withstand the cold and pressure differences


                  -Mudslide-like formations have been
                  found on Mars (left). These appear to
                  have been caused by water
                  movements. Psychrophiles may exist
                  there
Life in Outer Space?
                           -Europa is thought to have an ice
                           crust shielding a 30-mile deep
                           ocean. Reddish cracks (left) are
                           visible in the ice and may be
                           evidence of living populations


                           -Titan is enveloped with a hazy
                           gas (left) that is believed to
                           contain some organic
                           molecules, ie methane. This
                           may provide sustenance for life
                           on Titan’s surface

Images courtesy of the Current Science & Technology Center
Life in Outer Space?
                                  -Scientists have found that
                                  meteorites contain amino
                                  acids and simple sugars,
                                  very important building
                                  blocks. These may serve
                                  to spread life throughout
                                  the universe
Image courtesy of the Current Science & Technology Center


                                     -A sample of stratospheric
                                     air had shown a myriad of
                                     bacterial diversity 41 km
                                     above the earth’s surface
                                     (Lloyd, Harris, & Narlikar,
                                     2001)
                    Indeed, we may not be alone
A. The archaea are quite diverse, both in morphology and
   physiology
     1. They may stain gram positive or gram negative
     2. They may be spherical, rod-shaped, spiral, lobed,
        plate-shaped, irregularly shaped or pleomorphic
     3. They may exist as single cells, aggregates or filaments
     4. They may multiply by binary fission, budding,
        fragmentation, or other mechanisms
     5. They may be aerobic, facultatively anaerobic, or
        strictly anaerobic
     6. Nutritionally, they range from chemilithoautotrophs to
        organotrophs
     7. Some       are   mesophiles,     while    others    are
        hyperthermophiles that can grow above 100°C
     8. They are often found in extreme aquatic and terrestrial
        habitats; recently, archaea have been found in cold
        environments and may constitute up to 34% of the
        procaryotic biomass in Antarctic surface waters; a few
        are symbionts in animal digestive systems
Archaeal cell walls
  1. Archaea can stain either gram positive or gram negative,
     but their cell wall structure differs significantly from that of
     bacteria
        a. Many archaea that stain gram positive have a cell wall
           made of a single homogeneous layer
        b. The archaea that stain gram negative lack the outer
           membrane and complex peptidoglycan network
           associated with gram-negative bacteria
  2. Archaeal cell wall chemistry is different from that of
     bacteria
        a. Lacks muramic acid and D-amino acids and therefore
           is resistant to lysozyme and b-lactam antibiotics
        b. Some have pseudomurein, a peptidoglycan-like
           polymer that has L-amino acids in its cross-links and
           different monosaccharide subunits and linkage
        c. Others have different polysaccharides
  3. The archaea that stain gram negative have a layer of protein
     or glycoprotein outside their plasma membrane
Archaeal lipids and membranes
  1. Lipids have branched hydrocarbons attached to glycerol by
     ether links rather than straight-chain fatty acids attached to
     glycerol by ester links as seen in Bacteria and Eucarya
  2. Other, more complex tetraether structures are also found
  3. Membranes contain polar lipids such as phospholipids,
     sulfolipids, and glycolipids and also contain nonpolar lipids
     (7-30%), which are usually derivatives of squalene
  4. Membranes of extreme thermophiles are almost completely
     tetraether monolayers
F. • Archaeal Taxonomy-the new edition of Bergey’s Manual
     will divide the archaea into two phyla: Euryarchaeota and
     Crenarchaeota

• Phylum Crenarchaeota
  A. Many are extremely thermophilic, acidophilic, and sulfur-
     dependent
       1. Sulfur may be used as an electron acceptor in
          anaerobic respiration, or as an electron source by
          lithotrophs
       2. Almost all are strict anaerobes
       3. They grow in geothermally heated water or soils
          (solfatara) that contain elemental sulfur (sulfur-rich
          hot springs, waters surrounding submarine volcanic
          activity); some (e.g., Pyrodictum spp.) can grow quite
          well above the boiling point of water (optimum @
          105oC)
       4. Some are organotrophic; others are lithotrophic
       5. There are 69 genera; two of the better-studied genera
          are Sulfolobus and Thermoproteus
A. Sulfolobus
     1. Stain gram negative; are aerobic, irregularly lobed,
        spherical bacteria
     2. Thermoacidophiles
     3. Cell walls lack peptidoglycan but contain lipoproteins
        and carbohydrates
     4. Oxidize sulfur to sulfuric acid; oxygen is the normal
        electron acceptor, but ferric iron can also be used
     5. Sugars and amino acids may serve as carbon and
        energy sources
A. Thermoproteus
     1. Long, thin, bent or branched rods
     2. Cell wall is composed of glycoprotein
     3. Strict anaerobes
     4. They have temperature optima from 70-97°C and pH
        optima from 2.5 to 6.5
     5. They grow in hot springs and other hot aquatic
        habitats that contain elemental sulfur
     6. They carry out anaerobic respiration using organic
        molecules as electron donors and elemental sulfur as
        the electron acceptor; they can also grow
        lithotrophically using H2 and S0 as electron donors
        and CO or CO2 as the sole carbon source
• Phylum Euryarchaeota
  A. The Methanogens
       1. Strict anaerobes that obtain energy by converting CO2,
          H2, formate, methanol, acetate, and other compounds
          to either methane or to methane and CO2; there are at
          least five orders, which differ greatly in shape, 16S
          rRNA sequence, cell wall chemistry and structure,
          membrane lipids, and other features
       2. Methanogens belonging to the order Methanopyrales
          have been suggested to be among the earliest
          organisms to evolve on Earth
       3. Methanogenesis is an unusual metabolic process and
          methanogens contain several unique cofactors
       4. They thrive in anaerobic environments rich in organic
          matter, such as animal rumens and intestinal tracts,
          freshwater and marine sediments, swamps, marshes,
          hot springs, anaerobic sludge digesters, and even
          within anaerobic protozoa
       5. They are of great potential importance because
          methane is a clean-burning fuel and an excellent
          energy source
       6. They may be an ecological problem, however, because
          methane is a greenhouse gas that could contribute to
          global warming and also because methanogens can
          oxidize iron, which contributes significantly to the
          corrosion of iron pipes
A. The Halobacteria
     1. A group of extremely halophilic organisms divided
        into 15 genera
           a. They are aerobic chemoheterotrophs with
              respiratory metabolism; they require complex
              nutrients
           b. Motile or nonmotile by lophotrichous flagella
     2. They require at least 1.5 M NaCl and have growth
        optima near 3-4 M NaCl (if the NaCl concentration
        drops below 1.5 M the cell walls disintegrate; because
        of this they are found in high-salinity habitats and can
        cause spoilage of salted foods
     3. Halobacterium salinarum uses four different light-
        utilizing rhodopsin molecules
           a. Bacteriorhodopsin uses light energy to drive
              outward proton transport for ATP synthesis; thus
              they carry out a type of photosynthesis that does
              not involve chlorophyll
           b. Halorhodopsin uses light energy to transport
              chloride ions into the cell to maintain a 4-5 M
              intracellular KCl concentration
           c. Two other rhodopsins act as photoreceptors that
              control flagellar activity to position the bacterium
              in the water column at a location of high light
              intensity, but one in which the UV light is not
              sufficiently intense to be lethal
A. The Thermoplasms
0. Thermoacidic organisms that lack cell walls; only two
   genera are know: Thermoplasma and Picrophilus
     1. Thermoplasma
          a. Frequently found in coal mine refuse, in which
             chemolithotrophic bacteria oxidize iron pyrite to
             sulfuric acid and thereby produce a hot acidic
             environment
          b. Optimum temperature for growth of 55-59°C and
             an optimal PH of 1 to 2
          c. Cell membrane is strengthened by large
             quantities of diglycerol tetraethers,
             lipopolysaccharides, and glycoproteins
          d. Histonelike proteins stabilize their DNA; DNA-
             protein complex forms particles resembling
             eucaryotic nucleosomes
          e. At 59oC Thermoplasma takes the form of an
             irregular filament; the cells may be flagellated
             and motile
1. Picrophilus
                 a. Isolated from hot solfateric fields
                 b. Has an S-layer outside the plasma membrane
                 c. Irregularly shaped cocci with large cytoplasmic
                    cavities that are not membrane bounded
                 d. Aerobic and grows between 47°C and 65°C with
                    an optimum of 60°C
                 e. It grows only below pH 3.5, has an optimum of
                    pH 0.7 and will even grow at or near pH 0
A. Extremely thermophilic S0 metabolizers
0. Strictly anaerobic, reduce sulfur to sulfide
           1. Are motile by means of flagella
           2. Have optimum growth temperatures around 88-100°C
     B. Sulfate-reducing archaea
0. Gram-negative, irregular coccoid cells with walls of
   glycoprotein subunits
           1. Use a variety of electron donors (hydrogen, lactate,
              glucose) and reduce sulfite, sulfate, or thiosulfate to
              sulfide
           2. Are extremely thermophilic (optimum around 83°C);
              they are usually found near marine hydrothermal vents
           3. Contain two methanogen coenzymes
CONCLUSIONS
-Extremophiles are a very important and integral
  part of the earth’s biodiversity

They:
  - reveal much about the earth’s history and
  origins of life
  - possess amazing capabilities to survive in the
  extremes
  - are proving to be beneficial to both humans and
  the environment
  -may exist beyond earth

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Extremophiles imp. 1

  • 2. EXTREMOPHILES I. What are they? II. Types of Extremophiles III. Extreme Prokaryotes IV. Extreme Eukaryotes V. Extreme Viruses VI. Evolution of Extremophiles VII. Biotechnological Uses VIII. Industrial Uses IX. Extraterrestrial Extremophiles?
  • 3. What are Extremophiles? Extremophiles are microorganisms— whether viruses, prokaryotes, or eukaryotes— that survive under harsh environmental conditions that can include atypical temperature, pH, salinity, pressure, nutrient, oxic, water, and radiation levels
  • 4. Types of Extremophiles Types of Extremophiles
  • 5. Types of Extremophiles Other types include:  Barophiles -survive under high pressure levels, especially in deep sea vents  Osmophiles –survive in high sugar environments  Xerophiles -survive in hot deserts where water is scarce  Anaerobes -survive in habitats lacking oxygen  Microaerophiles -survive under low-oxygen conditions only  Endoliths –dwell in rocks and caves  Toxitolerants -organisms able to withstand high levels of damaging agents. For example, living in water saturated with benzene, or in the water-core of a nuclear reactor
  • 7. EXTREME PROKARYOTES Hyperthermophiles -Members of domains Bacteria and Archaea -Held by many scientists to have been the earliest organisms -Early earth was excessively hot, so these organisms would have been able to
  • 8. Morphology of Hyperthermophiles -Heat stable proteins that have more hydrophobic interiors, which prevents unfolding or denaturation at higher temperatures -Have chaperonin proteins that maintain folding -Monolayer membranes of dibiphytanyl tetraethers, consisting of saturated fatty acids which confer rigidity, preventing them from being degraded in high temperatures -Have a variety of DNA-preserving substances that reduce mutations and damage to nucleic acids, such as reverse DNA gyrase The red on these rocks and Sac7d is produced by -They can live without sunlight or organic Sulfolobus solfataricus, carbon as food, and instead survive on near Naples, Italy sulfur, hydrogen, and other materials that other organisms cannot metabolize
  • 9.
  • 11.
  • 12.
  • 13.
  • 14.
  • 15.
  • 16.
  • 17. a | Schematic representation of a cross-section of the cell envelope of Sulfolobus solfataricus showing the cytoplasmic membrane, with membrane-spanning tetraether lipids and an S-layer composed of two proteins — a surface-covering protein (red oval) and a membrane- anchoring protein (yellow oblong). b | Schematic representation of a cell envelope of an archaeon that stains positive with the Gram stain and that contains a pseudomurein layer in addition to the S-layer. The cytoplasmic membrane is composed of diether lipids.
  • 18.
  • 19.
  • 20.
  • 21.
  • 22.
  • 23.
  • 24. Some Hyperthermophiles Frequent habitats include volcanic vents and hot springs, as in the image to the left Pyrococcus abyssi 1µm Thermus aquaticus 1µm
  • 25. Deep Sea Extremophiles The deep-sea floor and hydrothermal vents involve the following conditions: low temperatures (2-3º C) – where only psychrophiles are present low nutrient levels – where only oligotrophs present high pressures – which increase at the rate of 1 atm for every 10 meters in depth (as we have learned, increased pressure leads to decreased enzyme- substrate binding) barotolerant microorganisms live at 1000-4000 meters A black smoker, a submarine barophilic microorganisms live at hot spring, which can reach depths greater than 4000 meters 518- 716°F (270-380°C)
  • 26. Extremophiles of Hydrothermal Vents Natural springs which vent warm or hot water on the sea floor near mid- ocean ridges. Associated with the spreading of 0.2µm 1µm the earth’s A cross-section of a bacterium A bacterial crust. High isolated from a vent. Often community from a temperatures such bacteria are filled with deep-sea and pressures viral particles which are hydrothermal vent abundant in hydrothermal near the Azores   vents
  • 27. Psychrophiles Some microorganisms thrive in temperatures well below the freezing point of water, such as in Some researchers believe that Antarctica psychrophiles live in conditions mirroring those found on Mars
  • 28. Psychrophiles possess: -proteins rich in α-helices and polar groups which allow for greater flexibility -“antifreeze proteins” that maintain liquid intracellular conditions by lowering freezing points of other biomolecules -membranes that are more fluid, containing unsaturated cis-fatty acids which help to prevent freezing -active transport at lower temperatures
  • 29. Halophiles -Divided into mild (1-6%NaCl), moderate (6-15%NaCl), and extreme (15-30%NaCl) -Halophiles are mostly obligate aerobic archaea How do halophiles survive high salt concentrations? -by interacting more strongly with water such as using more negatively charged amino acids in key structures -by making many small proteins inside the cell, and these, then, compete for the water -and by accumulating high levels of salt in the cell in order to outweigh the salt outside
  • 30. Barophiles -Survive under levels of pressure that are otherwise lethal to other organisms -Usually found deep in the earth, in the deep sea, hydrothermal vents, etc 1µm -scientists believe that A sample of barophilic barophiles may be bacteria from the able to survive on the earth’s interior Moon and other places in space
  • 31. Xerophiles Extremophiles which live in water- scarce habitats, such as deserts Produce desert varnish as seen in the image to the left Desert varnish is a thin coating of Mn, Fe, and clay on the surface of desert rocks, formed by colonies of bacteria living on the rock surface for thousands of years
  • 32. SOME COMMON GENERA OF PROKARYOTE EXTREMOPHILES 2um 1.8um 1um Thermotoga Aquifex Halobacterium 0.6um 0.9um 0.9um Methanosarcina Thermoplasma Thermococcus 1.3um 0.6um 0.7um Thermoproteus Pyrodictium Ignicoccus
  • 33. Deinococcus radiodurans The Radiation Resistor -Possesses extreme resistance to up to 4 million rad of radiation, genotoxic chemicals (those that harm DNA), oxidative damage from peroxides/superoxides, high levels of ionizing and ultraviolet radiation, and dehydration -It has from four to ten DNA molecules compared to only one for most other bacteria 0.8µm -Contains many DNA repair enzymes, such as RecA, which matches the shattered pieces of DNA and splices them back together. During these repairs, cell-building activities are shut off and the broken DNA pieces are kept in place
  • 34. Chroococcidiopsis The Cosmopolitan Extremophile 1.5µm -A cyanobacteria which can survive in a variety of harsh environments, such as hot springs, hypersaline habitats, hot, arid deserts throughout the world, and in the frigid Ross Desert in Antarctica -Possesses a variety of enzymes which assist in such adaptation
  • 35. Other Prokaryotic Extremophiles 1µm 1µm Gallionella ferrugineaand Anaerobic bacteria (iron bacteria), from a cave Current efforts in microbial taxonomy are isolating more and more previously undiscovered extremophile species, in places where life was least expected
  • 36. EXTREME EUKARYOTES THERMOPHILES/ACIDOPHILES 2µm
  • 37. EXTREME EUKARYOTES PSYCHROPHILES 2µm Snow Algae (Chlamydomonas nivalis) A bloom of Chloromonas rubroleosa in Antarctica These algae have successfully adapted to their harsh environment through the development of a number of adaptive features which include pigments to protect against high light, polyols (sugar alcohols, e.g. glycerine), sugars and lipids (oils), mucilage sheaths, motile stages and spore
  • 38. EXTREME EUKARYOTES ENDOLITHS Quartzite from Johnson Canyon, California. Sample shows green bands of endolithic algae. Rock is 9.5 cm wide -Endoliths (also called hypoliths) are usually algae, but can also be prokaryotic cyanobacteria, that exist within rocks and caves -Often are exposed to anoxic (no oxygen) and anhydric (no water) environments
  • 39. EXTREME EUKARYOTES PARASITES -Members of the Phylum Protozoa, which are regarded as the earliest eukaryotes to evolve, are mostly parasites -Parasitism is often a stressful relationship on both host and parasite, so they are considered extremophiles 20µm 15µm Trypanosoma gambiense, Balantidium coli, causes causes African sleeping dysentery-like symptoms sickness
  • 40. EXTREME VIRUSES Viruses are currently being isolated from habitats where temperatures exceed 200°F Instead of the usual icosahedral or rod-shaped capsids that known viruses possess, researchers have 40nm found viruses with novel Virus-like particles propeller-like structures isolated from the extreme environment of Yellowstone National Park These extreme viruses often hot springs live in hyperthermophile prokaryotes such as Sulfolobus
  • 41. CLASSIFICATION OF EXTREMOPHILES Phylogenetic Relationships Extremophiles are present among Bacteria, form the majority of Archaea, and also a few among the Eukarya
  • 42. PHYLOGENETIC RELATIONSHIPS -Members of Domain Bacteria (such as Aquifex and Thermotoga) that are closer to the root of the “tree of life” tend to be hyperthermophilic extremophiles -The Domain Archaea contain a multitude of extremophilic species: Phylum Euryarchaeota-consists of methanogens and extreme halophiles Phylum Crenarchaeota-consists of thermoacidophiles, which are extremophiles that live in hot, sulfur-rich, and acidic solfatara springs Phylum Korarchaeota-new phylum of yet uncultured archaea near the root of the Archaea branch, all are hyperthermophiles -Most extremophilic members of the Domain Eukarya are red and green algae
  • 44. The First Organisms? Early Earth was largely inhospitable: high CO 2/H2S/H2 etc, low oxygen, and high temperatures Lifeforms that could evolve in such an environment needed to adapt to these extreme conditions H2 was present in abundance in the early atmosphere. Many hyperthermophiles and archaea are H2 oxidizers Thus, it is widely held that extremophiles represent the earliest forms of life with non-extreme forms evolving after cyanobacteria had accumulated enough O2 in the atmosphere Results of rRNA and other molecular techniques have placed hyperthermophilic bacteria and archaea at the roots of the phylogenetic tree of life
  • 45. Evolutionary Theories  Consortia- symbiotic relationships between microorganisms, allows more than one species to exist in extreme habitats because one species provides nutrients to the others and vice versa  Genetic drift appears to have played a major role in how extremophiles evolved, with allele frequencies randomly changing in a microbial population. So alleles that conferred adaptation to harsh habitats increased in the population, giving rise to extremophile populations  Geographic isolation may also be a significant factor in extremophile evolution. Microorganisms that became isolated in more extreme areas may have evolved biochemical and morphological characteristics which enhanced survival as opposed to their relatives in more temperate areas. This involves genetic drift as well
  • 46. Slower Evolution -Extremophiles, especially hyperthermophiles, possess slow “evolutionary clocks” -That is, they have not evolved much from their ancestors as compared to other organisms -Hence, hyperthermophiles today are similar to hyperthermophiles of over 3 billion years ago -Slower evolution may be the direct result of living in extreme habitats and little competition -By contrast, other extremophiles, such as extreme halophiles and psychrophiles, appear to have undergone faster modes of evolution since they live in more specialized habitats that are not representative of early earth conditions
  • 47. Mat Consortia Mat Consortia A mat consortia in Yellowstone -Microbial mats consist of an upper layer of photosynthetic bacteria, with a lower layer of nonphotosynthetic bacteria -These consortia may explain some of the evolution that has taken place: extremophiles may have relied on other extremophiles and non-extremophiles for nutrients and shelter -Hence, evolution could have been cooperative
  • 48. USES OF EXTREMOPHILES HYPERTHERMOPHILES (SOURCE) USES DNA polymerases DNA amplification by PCR Alkaline phosphatase Diagnostics Proteases and lipases Dairy products Lipases, pullulanases and proteases Detergents Proteases Baking and brewing and amino acid production from keratin Amylases, α-glucosidase, pullulanase and xylose/glucose isomerases Baking and brewing and amino acid production from keratin Alcohol dehydrogenase Chemical synthesis Xylanases Paper bleaching Lenthionin Pharmaceutical S-layer proteins and lipids Molecular sieves Oil degrading microorganisms Surfactants for oil recovery Sulfur oxidizing microorganisms Bioleaching, coal & waste gas desulfurization Hyperthermophilic consortia Waste treatment and methane production
  • 49. USES OF EXTREMOPHILES PSYCHROPHILES (SOURCE) USES Alkaline phosphatase Molecular biology Proteases, lipases, cellulases and amylases Detergents Lipases and proteases Cheese manufacture and dairy production Proteases Contact-lens cleaning solutions, meat tenderizing Polyunsaturated fatty acids Food additives, dietary supplements Various enzymes Modifying flavors b-galactosidase Lactose hydrolysis in milk products Ice nucleating proteins Artificial snow, ice cream, other freezing applications in the food industry Ice minus microorganisms Frost protectants for sensitive plants Various enzymes (e.g. dehydrogenases) Biotransformations Various enzymes (e.g. oxidases)Bioremediation, environmental biosensors Methanogens Methane production
  • 50. USES OF EXTREMOPHILES HALOPHILES (SOURCE) USES Bacteriorhodopsin Optical switches and photocurrent generators in bioelectronics Polyhydroxyalkanoates Medical plastics Rheological polymers Oil recovery Eukaryotic homologues (e.g. myc oncogene product) Cancer detection, screening anti-tumor drugs Lipids Liposomes for drug delivery and cosmetic packaging Lipids Heating oil Compatible solutes Protein and cell protectants in variety of industrial uses, e.g. freezing, heating Various enzymes, e.g. nucleases, amylases, proteases Various industrial uses, e.g. flavoring agents g-linoleic acid, b-carotene and cell extracts, e.g. Spirulina and Dunaliella Health foods, dietary supplements, food coloring and feedstock Microorganisms Fermenting fish sauces and modifying food textures and flavors Microorganisms Waste transformation and degradation, e.g. hypersaline waste brines contaminated with a wide range of organics Membranes Surfactants for pharmaceuticals
  • 51. USES OF EXTREMOPHILES ALKALIPHILES (SOURCE) USES Proteases, cellulases, xylanases, lipases and pullulanases Detergents Proteases Gelatin removal on X-ray film Elastases, keritinases Hide dehairing Cyclodextrins Foodstuffs, chemicals and pharmaceuticals Xylanases and proteases Pulp bleaching Pectinases Fine papers, waste treatment and degumming Alkaliphilic halophiles Oil recovery Various microorganisms Antibiotics ACIDOPHILES (SOURCE) USES Sulfur oxidizing microorganisms Recovery of metals and desulfurication of coal Microorganisms Organic acids and solvents
  • 52. Taq Polymerase Isolated from the hyperthermophile Thermus aquaticus Much more heat stable Used as the DNA polymerase in the very useful Polymerase Chain Reaction (PCR) technique which amplifies DNA samples
  • 53. Alcohol Dehydrogenase -Alcohol dehydrogenase (ADH), is derived from a member of the archaea called Sulfolobus solfataricus -It works under some of nature's harshest volcanic conditions: It can survive to 88°C (190ºF) - nearly boiling - and corrosive acid conditions (pH=3.5) approaching the sulfuric acid found in a car battery (pH=2) -ADH catalyzes the conversion of alcohols and has considerable potential for biotechnology applications due to its stability under these extreme conditions
  • 54. Bacteriorhodopsin -Bacteriorhodopsin is a trans-membrane protein found in the cellular membrane of Halobacterium salinarium, which functions as a light- driven proton pump -Can be used for electrical generation
  • 55. Bioremediation - Bioremediation is the branch of biotechnology that uses biological processes to overcome environmental problems - Bioremediation is often used to degrade xenobiotics introduced into the environment through human error or negligence - Part of the cleanup effort after the 1989 Exxon Valdez oil spill included microorganisms induced to grow via nitrogen enrichment of the contaminated soil
  • 57. Psychrophiles as Bioremediators - Bioremediation applications with cold- adapted enzymes are being considered for the degradation of diesel oil and polychlorinated biphenyls (PCBs) - Health effects that have been associated with exposure to PCBs include acne-like skin conditions in adults and neurobehavioral and immunological changes in children. PCBs are known to cause cancer in animals
  • 58. An End to Pollution? New and innovative methods are being developed that utilize extremophiles for the elimination of pollution resulting from oil slicks, toxic chemical spills, derelict mines, etc
  • 59. Life in Outer Space? -Scientists have decided on 3 requirements for life: water energy carbon -Astrobiology: field of biology dealing with the existence of life beyond earth -Astrobiologists are currently looking for life on Mars, Jupiter’s moon Europa, and Saturn’s moon Titan -Such life is believed to consist of extremophiles that can withstand the cold and pressure differences -Mudslide-like formations have been found on Mars (left). These appear to have been caused by water movements. Psychrophiles may exist there
  • 60. Life in Outer Space? -Europa is thought to have an ice crust shielding a 30-mile deep ocean. Reddish cracks (left) are visible in the ice and may be evidence of living populations -Titan is enveloped with a hazy gas (left) that is believed to contain some organic molecules, ie methane. This may provide sustenance for life on Titan’s surface Images courtesy of the Current Science & Technology Center
  • 61. Life in Outer Space? -Scientists have found that meteorites contain amino acids and simple sugars, very important building blocks. These may serve to spread life throughout the universe Image courtesy of the Current Science & Technology Center -A sample of stratospheric air had shown a myriad of bacterial diversity 41 km above the earth’s surface (Lloyd, Harris, & Narlikar, 2001) Indeed, we may not be alone
  • 62. A. The archaea are quite diverse, both in morphology and physiology 1. They may stain gram positive or gram negative 2. They may be spherical, rod-shaped, spiral, lobed, plate-shaped, irregularly shaped or pleomorphic 3. They may exist as single cells, aggregates or filaments 4. They may multiply by binary fission, budding, fragmentation, or other mechanisms 5. They may be aerobic, facultatively anaerobic, or strictly anaerobic 6. Nutritionally, they range from chemilithoautotrophs to organotrophs 7. Some are mesophiles, while others are hyperthermophiles that can grow above 100°C 8. They are often found in extreme aquatic and terrestrial habitats; recently, archaea have been found in cold environments and may constitute up to 34% of the procaryotic biomass in Antarctic surface waters; a few are symbionts in animal digestive systems
  • 63. Archaeal cell walls 1. Archaea can stain either gram positive or gram negative, but their cell wall structure differs significantly from that of bacteria a. Many archaea that stain gram positive have a cell wall made of a single homogeneous layer b. The archaea that stain gram negative lack the outer membrane and complex peptidoglycan network associated with gram-negative bacteria 2. Archaeal cell wall chemistry is different from that of bacteria a. Lacks muramic acid and D-amino acids and therefore is resistant to lysozyme and b-lactam antibiotics b. Some have pseudomurein, a peptidoglycan-like polymer that has L-amino acids in its cross-links and different monosaccharide subunits and linkage c. Others have different polysaccharides 3. The archaea that stain gram negative have a layer of protein or glycoprotein outside their plasma membrane
  • 64. Archaeal lipids and membranes 1. Lipids have branched hydrocarbons attached to glycerol by ether links rather than straight-chain fatty acids attached to glycerol by ester links as seen in Bacteria and Eucarya 2. Other, more complex tetraether structures are also found 3. Membranes contain polar lipids such as phospholipids, sulfolipids, and glycolipids and also contain nonpolar lipids (7-30%), which are usually derivatives of squalene 4. Membranes of extreme thermophiles are almost completely tetraether monolayers
  • 65. F. • Archaeal Taxonomy-the new edition of Bergey’s Manual will divide the archaea into two phyla: Euryarchaeota and Crenarchaeota • Phylum Crenarchaeota A. Many are extremely thermophilic, acidophilic, and sulfur- dependent 1. Sulfur may be used as an electron acceptor in anaerobic respiration, or as an electron source by lithotrophs 2. Almost all are strict anaerobes 3. They grow in geothermally heated water or soils (solfatara) that contain elemental sulfur (sulfur-rich hot springs, waters surrounding submarine volcanic activity); some (e.g., Pyrodictum spp.) can grow quite well above the boiling point of water (optimum @ 105oC) 4. Some are organotrophic; others are lithotrophic 5. There are 69 genera; two of the better-studied genera are Sulfolobus and Thermoproteus
  • 66. A. Sulfolobus 1. Stain gram negative; are aerobic, irregularly lobed, spherical bacteria 2. Thermoacidophiles 3. Cell walls lack peptidoglycan but contain lipoproteins and carbohydrates 4. Oxidize sulfur to sulfuric acid; oxygen is the normal electron acceptor, but ferric iron can also be used 5. Sugars and amino acids may serve as carbon and energy sources
  • 67. A. Thermoproteus 1. Long, thin, bent or branched rods 2. Cell wall is composed of glycoprotein 3. Strict anaerobes 4. They have temperature optima from 70-97°C and pH optima from 2.5 to 6.5 5. They grow in hot springs and other hot aquatic habitats that contain elemental sulfur 6. They carry out anaerobic respiration using organic molecules as electron donors and elemental sulfur as the electron acceptor; they can also grow lithotrophically using H2 and S0 as electron donors and CO or CO2 as the sole carbon source
  • 68. • Phylum Euryarchaeota A. The Methanogens 1. Strict anaerobes that obtain energy by converting CO2, H2, formate, methanol, acetate, and other compounds to either methane or to methane and CO2; there are at least five orders, which differ greatly in shape, 16S rRNA sequence, cell wall chemistry and structure, membrane lipids, and other features 2. Methanogens belonging to the order Methanopyrales have been suggested to be among the earliest organisms to evolve on Earth 3. Methanogenesis is an unusual metabolic process and methanogens contain several unique cofactors 4. They thrive in anaerobic environments rich in organic matter, such as animal rumens and intestinal tracts, freshwater and marine sediments, swamps, marshes, hot springs, anaerobic sludge digesters, and even within anaerobic protozoa 5. They are of great potential importance because methane is a clean-burning fuel and an excellent energy source 6. They may be an ecological problem, however, because methane is a greenhouse gas that could contribute to global warming and also because methanogens can oxidize iron, which contributes significantly to the corrosion of iron pipes
  • 69. A. The Halobacteria 1. A group of extremely halophilic organisms divided into 15 genera a. They are aerobic chemoheterotrophs with respiratory metabolism; they require complex nutrients b. Motile or nonmotile by lophotrichous flagella 2. They require at least 1.5 M NaCl and have growth optima near 3-4 M NaCl (if the NaCl concentration drops below 1.5 M the cell walls disintegrate; because of this they are found in high-salinity habitats and can cause spoilage of salted foods 3. Halobacterium salinarum uses four different light- utilizing rhodopsin molecules a. Bacteriorhodopsin uses light energy to drive outward proton transport for ATP synthesis; thus they carry out a type of photosynthesis that does not involve chlorophyll b. Halorhodopsin uses light energy to transport chloride ions into the cell to maintain a 4-5 M intracellular KCl concentration c. Two other rhodopsins act as photoreceptors that control flagellar activity to position the bacterium in the water column at a location of high light intensity, but one in which the UV light is not sufficiently intense to be lethal
  • 70. A. The Thermoplasms 0. Thermoacidic organisms that lack cell walls; only two genera are know: Thermoplasma and Picrophilus 1. Thermoplasma a. Frequently found in coal mine refuse, in which chemolithotrophic bacteria oxidize iron pyrite to sulfuric acid and thereby produce a hot acidic environment b. Optimum temperature for growth of 55-59°C and an optimal PH of 1 to 2 c. Cell membrane is strengthened by large quantities of diglycerol tetraethers, lipopolysaccharides, and glycoproteins d. Histonelike proteins stabilize their DNA; DNA- protein complex forms particles resembling eucaryotic nucleosomes e. At 59oC Thermoplasma takes the form of an irregular filament; the cells may be flagellated and motile
  • 71. 1. Picrophilus a. Isolated from hot solfateric fields b. Has an S-layer outside the plasma membrane c. Irregularly shaped cocci with large cytoplasmic cavities that are not membrane bounded d. Aerobic and grows between 47°C and 65°C with an optimum of 60°C e. It grows only below pH 3.5, has an optimum of pH 0.7 and will even grow at or near pH 0
  • 72. A. Extremely thermophilic S0 metabolizers 0. Strictly anaerobic, reduce sulfur to sulfide 1. Are motile by means of flagella 2. Have optimum growth temperatures around 88-100°C B. Sulfate-reducing archaea 0. Gram-negative, irregular coccoid cells with walls of glycoprotein subunits 1. Use a variety of electron donors (hydrogen, lactate, glucose) and reduce sulfite, sulfate, or thiosulfate to sulfide 2. Are extremely thermophilic (optimum around 83°C); they are usually found near marine hydrothermal vents 3. Contain two methanogen coenzymes
  • 73. CONCLUSIONS -Extremophiles are a very important and integral part of the earth’s biodiversity They: - reveal much about the earth’s history and origins of life - possess amazing capabilities to survive in the extremes - are proving to be beneficial to both humans and the environment -may exist beyond earth

Hinweis der Redaktion

  1. Figure: 04-34 Caption: Pseudopeptidoglycan and S-layers. (a) Structure of pseudopeptidoglycan, the cell wall polymer of Methanobacterium species. Note the resemblance to the structure of peptidoglycan shown in Figure 4.30, especially the peptide cross-links, in this case between N -acetyltalosaminuronic acid (NAT) residues instead of muramic acid residues. NAG, N -Acetylglucosamine.