Extreme Biology
Energy
• Living things use food as a source of energy
– Autotrophs make their own food.
Autotrophs capture the sun’s energy and use it along with water and carbon dioxide to make their own food.
• Auto (self)
• Troph ( feeder)
– Heterotrphs cannot make their own food. They get energy from the sun indirectly by eating autotrophs or other heterotrophs
• Hetero (other)
• Troph (feeder)
Universal Tree of Life: 3 Domain System
Bacteria and Archaea are both prokaryotes
Extreme Environments on Earth
1. Sea Ice (extreme cold)
2. Hydrothermal vents (extreme heat and high metal content) 3. Sulfuric Springs (extreme heat and highly acidic)
4. Salt Lake (extreme salt concentrations)
5. Soda Lake (extreme salt concentration and highly alkaline)
Cellular Targets of Adaptations to Extreme Environments
Cytoplasm: water, proteins, metabolites, salts
Nucleoid: Aggregated DNA Chromosome
Typically lipid bilayer
Typical Prokaryotic Cell
Life on Ice
Over 75% of Earth’s biosphere is permanently cold (< 5°C)
Much of the life present in the cold environs is planktonic growth of bacteria and archaea in frigid marine waters (~104 cells/ml) (psychrophiles)
Identified using rRNA techniques – 16S rRNA sequencing
– Fluorescent rRNA DNA probes
At this point physiology of psychrophilic archaea/bacteria undetermined
Cold adaptations: more fluid membranes, more structurally flexible proteins
Psychrophilic cyanobacteria
Methanogenium frigidum
Adaptations to Extreme Cold: Making More Fluid Membranes
More fluid membranes result from putting unsaturated/polyunsaturated fatty acids into the membrane
More Life on Ice: Algae
Algae living on the ice
(photosynthetic unicellular plant)
Lichen = symbiotic relationship between algae and fungi
Phytoplankton Krill
Polychaete Worms Living on Methane Ice
It is thought that the worms eat the bacteria that are growing on the methane ice
Lake Vostoc: A model for Life on Europa?
Hydrothermal Vent Systems
Anatomy of A Vent
Hydrothermal Vents: Abiotic Conditions
Extremely hot temperatures (> 350ºC [hydrostatic pressure of 265 atm prevents water from boiling until 460 ºC ])
Extremely high pressures up to 1,000 atm
Vents rich in minerals (eg. Iron oxides, sulfates, sulfides, manganese oxides, calcium, zinc, and copper sulfides)
Hot waters anaerobic since solubility of oxygen decreases as water temperature increases
Hydrothermal Vents: Biotic Community
Archaea and bacteria grow in or near vent chimneys, shown to live and reproduce at temp. of 115°C (hyperthermophiles)
As of 5 years ago believed highest upper temp. for life was 105
°C, now expect hyperthermophiles may grow up to 160 °C [limit of ATP stability]
Rich microbial communities grow at some distance from vent chimneys where temperatures are more moderate (8 - 12°C) due to mixing mixing with cold seawater (~2°C)
Hydrothermal Vent Ecosystems: The Prokaryotes
Methanococcus janaschii (85°C)
Pyrococcus furiosus (100°C)
Vent contact slide Aquifex aeolicus (95°C)
Thermotoga maritima (90°C)
Archaea Bacteria
Archaeoglobus fulgidus (83°C)
Thermal Adaptations Used By Hyperthermophiles for Survival
Membrane: ether-linked membrane-lipids, monolayer membranes
Protein: hydrophobic protein core, salt bridges, chaperonins
DNA: Cation stabilization (Mg2+), Reverse DNA gyrase, DNA-Binding proteins (histones)
General: compatible solutes?
Histone and DNA
Hydrothermal Vent Ecosystem: Tube Worms
Vent water is ~350o C with high H2S concentrations
Surrounding water is ~10-20oC
Gutless tubeworms (Riftia have a mutualistic symbiosis with aerobic H2S- oxidizing bacteria (Thiomicrospira).
Vestimentiferan worms; Riftia pachyptile
Endosymbiosis in Tubeworms
Hydrothermal Vent Ecosystems: Bivalves
Calyptogena magnifica Bathymodiolus thermophilus
Hydrothermal Vent Ecosystems: “Snow Flurries” and Crabs
Flocs of sulfur bacteria Galatheid crabs
And Where There’s Crabs, Octopi Are Not Far Behind
Black Smokers – Sulfur Reducers
• Black smoker vents
• Found in deepest parts of the ocean
• Volcanic, mineral-enriched water outflows
• Rich in iron, sulfur compounds
• Very little/no oxygen
• Discovered in the 1970s
• Temps as high as 750 F (!!)
• Does not boil, though, due to extreme pressure at this depth
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Black Smoker Structure
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Black Smoker Ecology
• Deep sea exploration vehicles investigate black smokers in the 1980’s
• Much to everyone’s surprise, they find LIFE !!
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Black Smoker Ecology
• Not just life – fully-developed ecosystems!
• Crabs, shrimp, clams, Pompeii worms
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Pompeii Worms
• Tube worms anchored near black smoker vents
• Bottom end has very high temps; top end more like 70F
• Hot water flows through tubes; length as much as 10 feet!
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Pompeii Worms
• “Hairy” back is heat-resistant microbe mat (symbiotic with worm mucus)
• Red “feathers” include hemoglobin; separates hydrogen sulfide from vent flow
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Extreme Environments on Earth
1. Sea Ice (extreme cold)
2. Hydrothermal vents (extreme heat and high metal content) 3. Sulfuric Springs (extreme heat and highly acidic)
4. Salt Lake (extreme salt concentrations)
5. Soda Lake (extreme salt concentration and highly alkaline)
Life in Sulfur Springs (Hot and Acidic)
Abiotic conditions:
- high temperatures >30°C - low pH (< 4)
- high sulfur
Sulfur-oxidizing, acid-loving, hyperthermophiles such as the archaeon Sulfolobus have been isolated from sulfur hot springs
Sulfolobus grows at 90oC, pH 1-5
–Oxidizes H2S (or So) to H2SO4 –Fixes CO2 as sole C-source
Acidophiles do not have low internal pH’s and have adapted to keep protons outside the cell
Other Acidic Environments and Denizens
Acid mine drainage Acidophilic archaeon, Picrophilus oshimae, grows optimally at pH 0.7, cannot grow above pH 4
Red alga Cyanidarium caldarium grows at pH of 0.5
Archaeaon Ferroplasma acidarmanus thrives in acid mine drainage at pH 0 (has no cell wall)
Acidophiles studied to date appear to have very efficient membrane-bound Na+/H+ pumps and membranes with low permeability to protons
High Salt Environments
Salt evaporation ponds
Great Salt Lake
Low biodiversity; only home to halophilic organisms belonging to Archaea, Bacteria and some algae
Extreme halophiles require at least 1.5 M NaCl for growth (most need 2 – 4 M NaCl for optimum growth)
Cell lysis occurs below 1.5 M
Membranes are stabilized by Na+
Maintain high internal K+Cl- to balance high external Na+Cl-
A number of halophiles have a unique type of
“photosynthesis”
Multiple light-sensitive proteins
–Halorhodopsin (Cl- transport, creating Cl- gradient which drives K+ uptake)
–Bacteriorhodopsin (photosynthesis?)
Halophilic Algae
Dunaliella salina
Photosynthetic flagellate
Red because of high
concentrations of beta-carotene
On sensing high salinity, pumps out Na+ ions and replaces with K+ ions
In high salt, will alter
photosynthetic pathway to produce glycerol (water-soluble, nonionic substance which prevents
dehydration) instead of starch
Halobacterium salinarum and Light-mediated ATP Synthesis
Halobacterium salinarum
Halobacterium contain photopigments which are used to synthesize ATP as a result of proton motive force generation
cis-form
trans-form
light
Retinal chromophore of bacteriorhodopsin
High Salt Alkaline Environments: Soda Lakes
Lake Magadi (Soda lake in Kenya)
Have very high pH (> 9) due to high levels of CO32- ion
Very few organisms can tolerate alkaline conditions (to date only alkalophilic prokaryotes have been isolated)
Most alkalophilic organism,
cyanobacterium Plectonema, grows at pH of 13
Alkalophile adaptations: pumps to pump out OH-, efficient Na+/H+ to provide internal H+, modified
membranes
Cyanobacterium
Spirilina Natronobacterium
Survival Under Conditions of High Level Radiation Exposure: Deinococcus radiodurans
Aerobic, mesophilic bacterium
Extremely resistant to desiccation, UV and ionizing radiation -- Can survive 3-5 million rads (100 rads is lethal for humans)
Contain variable numbers (4-10) of chromosomes
DNA Damage Repair in Deinococcus radiodurans
Deinococcus radiodurans has very efficient DNA repair machinery
DNA sheared by radiation will
reform within 24h
Importance of Extremophiles:Extremozymes
Enzymes from extremophiles offer some important potential benefits:
Hyperthermophiles
– Sugar conversions without microbial growth and contamination
Psychrophiles
– Modification of flavor/texture of foods without microbial growth & spoilage
Acidophiles
– Removal of sulfur from coal & oil
Alkalophiles
– Cellulases that can be used in detergents
Importance of Extremophiles: Astrobiological Implications
Mars
Europa
Extreme environments on Earth are thought to be very similar to extreme
environments that exist elsewhere in space
Microorganisms that thrive in Earth extreme environments are thought to be likely candidates for the types of biota that may exist in extraterrestrial habitats
Mars is postulated to have extremophilic regions including permafrost,
hydrothermal vents, and evaporite crystals
Europa is thought to have a subsurface ocean
Extreme Life: Aquifex Aeolicus
• In the 1960’s, biologists
were interested in studying
“how extreme” life could be
• They knew that microbes lived in water downstream from hot springs in
Yellowstone National Park
• The springs themselves reached temperatures of
~85C (185 F) – near the boiling point of water
• The question: How far upstream (close to the hottest water) could microbes survive?
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Aquifex Aeolicus Surprise
• Biologists discovered bacteria in the hottest parts of the hot springs themselves
• These creatures survive – even thrive and
reproduce!! – at ~85C (185 F), near the boiling point of water
• Picture shows microbial mats (as in stromatolites) in Yellowstone hot spring
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Aquifex Aeolicus Properties
• These are very small bacteria
• Prokaryotes
• Genome structure is only 1/3 as long (complex) as E. coli (a model “simple” bacteria)
• Single DNA molecule in a circular chromosome
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Aquifex Aeolicus Metabolism
• A. aeolicus survives from H, O, CO2, and mineral salts
• Requires oxygen for
respiration (so, not that primitive)
• But … no need for sunlight, nor sunlight-using food !!
• Purely chemical food source (in the presence of thermal energy from the water)
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The colors of Prismatic Spring in
Yellowstone come primarily from the hyperthermophile microbes in it