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Archaea
17.1 Phylogenetic Overview of Archaea 17.2 Energy Conservation and Autotrophy in Archaea
Archaea share many characteristics with both Bacteria and Eukarya Archaea are split into two major groups
Crenarchaeota
Euryarchaeota
Figure 17.1
- Glucose metabolism
: EMP or slightly modified Entner-Doudoroff pathway
fumarate)
: establish proton motive force : ATP synthesis through membrane-bound ATPase
- Chemolithotrophy
: H2 as a common electron donor and energy source is well established
most hyperthermophiles
Reverse TCA cycle in some hyperthermophiles Calvin cycle in Methanococcus jannaschii and a Pyrococcus species (both are hyperthermophiles)
II. Euryarchaeota
17.3 Extremely Halophilic Archaea 17.4 Methane-Producing Archaea: Methanogens 17.5 Thermoplasmatales
II. Euryarchaeota
Euryarchaeota
Physiologically diverse group of Archaea Many inhabit extreme environments
E.g., high temperature, high salt, high acid
Haloarchaea
Extremely halophilic Archaea Have a requirement for high salt concentrations
Typically require at least 1.5 M (~9%) NaCl for growth
Found in solar salt evaporation ponds, salt lakes, and artificial saline habitats (i.e., salted foods)
Figure 17.2a
Figure 17.2b
Figure 17.2c
Figure 17.2d
Salt lakes can vary in ionic composition, selecting for different microbes
Great Salt Lake similar to concentrated seawater Soda lakes are highly alkaline hypersaline environments
Haloarchaea
Reproduce by binary fission Do not form resting stages or spores
Dividing cell
Figure 17.3
Halobacterium species instead pump large amounts of K+ into the cell from the environment
Intracellular K+ concentration exceeds extracellular Na+ concentration and positive water balance is maintained
Proteins of halophiles
Highly acidic Contain fewer hydrophobic amino acids and lysine
residues
Figure 17.4
Sensory rhodopsins
Control phototaxis
Figure 17.5
Habitats of Methanogens
Methanobrevibacter ruminantium
Figure 17.6a
Methanobrevibacter arboriphilus
Figure 17.6b
Methanospirillum hungatei
Figure 17.6c
Methanosarcina barkeri
Figure 17.6d
Diversity of Methanogens
Demonstrate diversity of cell wall chemistries
Pseudomurein (e.g., Methanobacterium,
Methanobrevibacter)
Methanochondroitin (e.g., Methanosarcina)
- (N-acetylgalactosamine + glucuronic acid)n
Methanobrevibacter ruminantium
Methanosarcina barkeri
Figure 17.7a
Methanocaldococcus jannaschii
Figure 17.8a
17.5 Thermoplasmatales
Methanocaldococcus jannaschii as a model methanogen
Contains 1.66 mB circular genome with about 1,700 genes Genes for central metabolic pathways and cell division
- Similar to those in Bacteria
Thermoplasmatales
Taxonomic order within the Euryarchaeota
Contains 3 genera
Thermoplasma Ferroplasma
Picrophilus
Thermophilic and/or extremely acidophilic Thermoplasma and Ferroplasma lack cell walls
Thermoplasma
Chemoorganotrophs Facultative aerobes via sulfur respiration
Thermophilic
Acidophilic
Thermoplasma Species
Thermoplasma acidophilum
Figure 17.9a
Thermoplasma Species
Figure 17.9b
Figure 17.10
Thermoplasma (contd)
Evolved unique cytoplasmic membrane structure to maintain positive osmotic pressure and tolerate high temperatures and low pHs
Membrane contains lipopolysaccharide-like material (lipoglycan) consisting of tetraether lipid monolayer membrane with mannose and glucose Membrane also contains glycoproteins but not sterols
Figure 17.11
Ferroplasma
Chemolithotrophic Acidophilic Oxidizes Fe2+ to Fe3+, uses CO2 as carbon source Grows in mine tailings containing pyrite (FeS)
- Generates acid (acid mine drainage)
Picrophilus
Extreme acidophiles
Grow optimally at pH 0.7
Have an S-layer
Model microbe for extreme acid tolerance
Pyrococcus
Methanopyrus
Figure 17.1
Thermococcales
Distinct order that contains Thermococcus and Pyrococcus
Thermococcus: organics + So, 55-95oC Pyrococcus: organics + So, opt. 100oC, 70-106oC
Figure 17.12b
Methanopyrus
Methanogenic (CO2 + H2)
Methanopyrus
Figure 17.13a
Methanopyrus
Figure 17.13b
17.7 Archaeoglobales
Archaeoglobales
Hyperthermophilic Couple oxidation of H2, lactate, pyruvate, glucose, or complex organic compounds to the reduction of SO42- to H2S
Archaeoglobus
Opt. temp. 83oC Produce methane, but lacks genes for methyl-CoM reductase
Ferroglobus
Opt. temp. 85oC Fe2+ + NO3- Fe3+ + NO2- + NO
Archaeoglobales
Figure 17.14a
Figure 17.14b
Nanorchaeum equitans
One of the smallest cellular organisms (~0.4 m) Obligate symbiont of the crenarchaeote Ignicoccus
Nanoarchaeum
Ignicoccus Nanoarchaeum
Figure 17.15a
Figure 17.15b
Aciduliprofundum
Thermophilic: 55-75oC Acidophile: pH 3.3-5.8, lives in sulfide deposits in hydrothermal vents Oragnics + So or Fe3+
III. Crenarchaeota
17.9 Habitats and Energy Metabolism of Crenarchaeota 17.10 Hyperthermophiles from Terrestrial Volcanic Habitats 17.11 Hyperthermophiles from Submarine Volcanic
Habitats
17.12 Nonthermophilic Crenarchaeota
Crenarchaeota
Inhabit temperature extremes Most cultured representatives are hyperthermophiles
Habitats of Crenarchaeota
Figure 17.16a
Figure 17.16b
Figure 17.16c
Figure 17.16d
Hyperthermophilic Crenarchaeota
Most are obligate anaerobes Chemoorganotrophs or chemolithotrophs with diverse
Acidianus
Also lives in acidic sulfur hot springs Able to grow using elemental sulfur both aerobically and anaerobically (as an electron donor and electron acceptor, respectively)
Sulfolobus acidocaldarius
Figure 17.17a
Acidianus infernus
Figure 17.17b
Thermoproteales
An order containing the key genera Thermoproteus, Thermofilum, and Pyrobaculum
Thermoproteus neutrophilus
Figure 17.18a
Thermofilum librum
Figure 17.18b
Pyrobaculum aerophilum
Figure 17.18c
Staphylothermus
Pyrodictium occultum
Figure 17.19a
Figure 17.19c
Negative stain of a cell of strain 121, the most heat-loving of all known
Figure 17.19d
Figure 17.20a
Figure 17.20b
Figure 17.21
Cold-Dwelling Crenarchaeota
DAPI (diamidino-2-phenylindole) stained
17.13 An Upper Temperature Limit for Microbial Life 17.14 Adaptations to Life at High Temperature 17.15 Hyperthermophilic Archaea, H2, and Microbial
Evolution
Figure 17.23
solutes
Use of more heat-stable molecules
e.g., use of nonheme iron proteins instead of proteins that use NAD and NADH
Chaperonins
Class of proteins that refold partially denatured proteins Thermosome
A major chaperonin protein complex in Pyrodictium
DNA Stability
High intracellular solute levels stabilize DNA
Reverse DNA gyrase
Introduces positive supercoils into DNA, which stabilizes DNA Found only in hyperthermophiles
High intracellular levels of polyamines (e.g., putrescine, spermidine) stabilize DNA and RNA DNA-binding proteins (archaeal histones) compact DNA into nucleosome-like structures
Figure 17.25
Lipid Stability
Possess dibiphytanyl tetraether type lipids; form a lipid
Figure 17.26