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The Archaeal Domain Methanogens and the C Cycle Estimated global production of methane 109 tons/yr. A cow can produce 100 liters of methane a day. Methane is an important greenhouse gas. Methanogens are found in many places in the Euryarchaeota. Methanogens Thermophilic species H2 and CO2 to make CH4. Methanocaldococcus jannaschii - 85˚C Methanopyrus kandleri - 100˚C Methanopyrus: -isolated from sediments near submarine hydrothermal vent chimney -generation time is 1hr at 100˚C -branches at the base of the archaeal tree Mesophilic species can also make methane from simple organic compounds (formate, acetate, methanol, methylamines) Euryarchaeota Thermoplasma: thermoacidophile aerobic or anaerobic sulfur respiration found in acidic soils and coal refuse piles Picrophilus: related to Thermoplasma grows optimally at pH 0.6 (can grow at pH -0.06!) membranes leak at pH 4 solfataras Early branching hydrothermal Vent Euryarchaeota. Thermococcus (“hot ball”, growing at 70-95˚C): spherical, highly motile anaerobic chemoorganotroph Pyrococcus (“fire ball”, growing at 70-106˚C): close relative of Thermococcus Thermococcus celer Pyrococcus furiosus Archaeoglobus Hyperthermophilic sulfate reducer. Hot marine sediments and hydrothermal vents. Shared many unique traits with methanogens: weird enzymes Cultures produce small amounts of methane. Closely related to methanogens. Sulfate reduction genes from the bacterial domain via lateral gene transfer. . Halobacterium, Haloferax, Natronobacterium Late branches in Euryarchaeota. Aerobic organotrophs. Halo- in neutral pH environments. Nat- in alkaline environments. Halobacterium Halogeometricum Haloarcula Natronococcus Crenarchaeota QuickT ime™ and a TI FF (Uncompressed) decompressor are needed to see this picture. Sulfolobus: S-rich acidic hot springs thermoacidophile aerobic chemotroph oxidizes H2S or S˚ to H2SO4 Sulfur Caldron Crenarchaeota, cont. Thermoproteus: long thin rods strict anaerobes S˚ reducer (likely an ancient metabolism) Submarine Vent Crenarchaeota Pyrodictium (“fire net”): Topt 105˚C network of fibers attach to other cells strict anaerobe chemolithotroph or chemoorganotroph Pyrolobus (“fire lobe”): Topt 106˚C holds the upper temperature record (species can grow > 113˚C) walls of black smoker chimney chemoautotroph Summary Euryarchaeota: methanogens Archaeoglobus thermoacidophiles - sulfur respiration halophiles Crenarchaeota many hyperthermophiles many organotrophs sulfur respiration acidophiles and neutrophiles Lecture 20. Proterozoic Earth and the Rise in Oxygen reading: Chapter 4 Hadean oldest rocks on Earth - end of heavy bombardment origin of the solar system Hadean billions of years ago: 4.56 rise in oxygen first multiplate tectonics? cellular fossils Archean 3.8 Proterozoic 2.5 Heavy bombardment. Delivery of volatiles. Possible early oceans. Warm early Earth but a faint young Sun. Few rocks. Cambrian Explosion Phanerozoic 0.55 present Archean QuickTime™ and a TIFF (Uncompressed) decompressor are needed to see this picture. Stromatolites and banded iron formation, particularly at end of Archean. Greenstone belts giving rise to continents. Creation of continental shields. Great carbonate reefs. Beginnings of life. Proterozoic origin of the solar system oldest rocks on Earth - end of heavy bombardment Hadean billions of years ago: 4.56 rise in oxygen first multiplate tectonics? cellular fossils Archean 3.8 Split into 3 eras: Paleoproterozoic Mesoproterozoic Neoproterozoic Proterozoic 2.5 3.8/3.5-1.6 Ga 1.6-0.9 Ga 900-543 Ma Cambrian Explosion Phanerozoic 0.55 present Paleoproterozoic, 2.8-1.6 Ga A Time of Fundamental Transitions: 2.4 Ga sulfur isotopic signatures of sulfate reducing bacteria appear 2.32 Ga oxygenation of the atmosphere 2.3 Ga global glaciations 2.2-2.1 Ga disruptions in carbon isotopes 1.8 Ga banded iron formation disappears Sulfur Isotopes microbes prefer light isotope of sulfur 32S over 33S & 34S sulfate reducers: 2”CH2O” + SO422H2S + Fe2+ -----> FeS2 ---> 2HCO3- + H2S + 2H2 pyrite in sedimentary rocks records presence of sulfate reducers <--- sediments enriched in 34S <--- sediments enriched in 32S Stepwise Oxygenation of the Earth Stage I: Before 2.32 Ga O2 < 10-5 present atmospheric levels (PAL) Very low levels of oxygen. Stratified oceans. Carbonates on the surface, iron rich at depth. Stage 2: Transition period 2.32 - 1.8 Ga Intermediate levels of oxygen. Oceans still stratified. Carbonates and oxides on the surface, iron rich at depth. Stage 3: O2 rises to levels similar to what is seen today. Oceans no longer stratified - similar to today. Global Glaciations Evidence of: massive glaciers worldwide near the equator ~2.3 Ga Disruptions in the Carbon Isotopes Disruptions in the Carbon Isotopes, cont. “Odd” carbon isotopes recorded in: organic material - highly enriched in 12C and in carbonate rocks highly enriched in 13C Don’t know the cause. Global disruption in the carbon cycle? Disappearance of BIF Lots of BIF deposited in the Paleoproterozoic. But by 1.8 Ga it disappears! QuickTime™ and a TIFF (Uncompressed) decompressor are needed to see this picture. Oceans sediments no longer contain iron oxides, but contain iron sulfides (like pyrite) increasing S content of the oceans. QuickTime™ and a TIFF (Uncompressed) decompressor are needed to see this picture. Appearance of Familiar Microbial Fossils Large diameter microfossils thought to be cyanobacteria appear at ~2.15 Ga. By 1.8 Ga, fossil akinetes - produced by one group of cyanobacteria are seen. this image is copyrighted by the Precambrian Paleobiology Research Group - UCLA Lecture 21. Basic Architecture of the Eukaryotic Cell, Symbioses, Early Eukaryote Fossils. reading: Chapter 5