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19.1 Phylogenetic and Metabolic Diversity of Archaea Archaea share many characteristics with both Bacteria and Eukarya Archaea are split into two major groups (Figure 19.1) Crenarchaeota Euryarchaeota © 2012 Pearson Education, Inc.

19.1 Phylogenetic and Metabolic Diversity of Archaea Archaea share many characteristics with both Bacteria and Eukarya Archaea are split into two major

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19.1 Phylogenetic and Metabolic Diversity of Archaea

• Archaea share many characteristics with both Bacteria and Eukarya

• Archaea are split into two major groups (Figure 19.1)– Crenarchaeota

– Euryarchaeota

© 2012 Pearson Education, Inc.

19.1 Phylogenetic and Metabolic Diversity of Archaea

• Bioenergetics and intermediary metabolism of Archaea are similar to those found in Bacteria– Except some Archaea use methanogenesis

– Autotrophy via several different pathways is widespread in Archaea

© 2012 Pearson Education, Inc.

Figure 19.1

Extreme halophiles

Extremeacidophiles

Hyperthermophiles

Crenarchaeota

Marine Crenarchaeota

Pyrodictium

Thermoproteus

Sulfolobus

Desulfurococcus

EuryarchaeotaMarine Euryarchaeota

Halophilicmethanogen

Halobacterium

Halococcus

Natronococcus

Archaeoglobus

Methanobacterium Methanocaldococcus

Methanothermus

Thermococcus/Pyrococcus

Nanoarchaeum

Methanopyrus

Methanosarcina

Methanospirillum

Thermoplasma

Ferroplasma

Picrophilus

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II. Euryarchaeota

• Euryarchaeota– Physiologically diverse group of Archaea

– Many inhabit extreme environments • Examples: high temperature, high salt, high acid

© 2012 Pearson Education, Inc.

19.2 Extremely Halophilic Archaea

• Haloarchaea• Key genera: Halobacterium, Haloferax,

Natronobacterium– Extremely halophilic Archaea

– Have a requirement for high salt concentrations

• Typically require at least 1.5 M (~9%) NaCl for growth

– Found in artificial saline habitats (e.g., salted foods), solar salt evaporation ponds, and salt lakes (Figure 19.2)

© 2012 Pearson Education, Inc.

Figure 19.2

© 2012 Pearson Education, Inc.

19.2 Extremely Halophilic Archaea

• Haloarchaea– Reproduce by binary fission

– Do not form resting stages or spores

– Most are nonmotile

– Most are obligate aerobes

– Possess adaptations to life in highly ionic environments

• Cell wall is composed of glycoprotein and stabilized by Na+ (Figure 19.3)

© 2012 Pearson Education, Inc.

19.2 Extremely Halophilic Archaea

• Water Balance in Extreme Halophiles– Halophiles need to maintain osmotic balance

• This is usually achieved by accumulation or synthesis of compatible solutes

– 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

© 2012 Pearson Education, Inc.

19.2 Extremely Halophilic Archaea

• Proteins of halophiles– Are highly acidic

– Contain fewer hydrophobic amino acids and lysine residues

© 2012 Pearson Education, Inc.

19.2 Extremely Halophilic Archaea

• Some haloarchaea are capable of light-driven synthesis of ATP (Figure 19.4)

– Bacteriorhodopsin• Cytoplasmic membrane proteins that can

absorb light energy and pump protons across the membrane

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Animation: Bacteriorhodopsin and Light Mediated ATP SynthesisAnimation: Bacteriorhodopsin and Light Mediated ATP Synthesis

Figure 19.4

Out In

Membrane

Bacteriorhodopsin

ATPase

© 2012 Pearson Education, Inc.

19.3 Methanogenic Archaea

• Methanogens (Figure 19.5)– Key genera: Methanobacterium,

Methanocaldococcus, Methanosarcina• Microbes that produce CH4

• Found in many diverse environments• Taxonomy based on phenotypic and phylogenetic

features

• Process of methanogenesis first demonstrated over 200 years ago by Alessandro Volta

© 2012 Pearson Education, Inc.

Figure 19.5

© 2012 Pearson Education, Inc.

19.3 Methanogenic Archaea

• Diversity of Methanogens– Demonstrate diversity of cell wall chemistries

(Figure 19.6 and Figure 19.7)• Pseudomurein (e.g., Methanobacterium)• Methanochondroitin (e.g., Methanosarcina)• Protein or glycoprotein (e.g.,

Methanocaldococcus)• S-layers (e.g., Methanospirillum)

© 2012 Pearson Education, Inc.

Figure 19.6

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Figure 19.7

© 2012 Pearson Education, Inc.

19.5 Thermococcales and Methanopyrus

• Three phylogenetically related genera of hyperthermophilic Euryarchaeota:

– Thermococcus

– Pyrococcus

– Methanopyrus

• Comprise a branch near root of archaeal tree

© 2012 Pearson Education, Inc.

19.5 Thermococcales and Methanopyrus

• Thermococcales– Distinct order that contains Thermococcus

and Pyrococcus (Figure 19.11)

– Indigenous to anoxic thermal waters

– Highly motile

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Figure 19.11

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Figure 19.16

© 2012 Pearson Education, Inc.