The Tree of Life. Metabolic Pathways. Calculation Of Energy Yields

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1 The Tree of Life Metabolic Pathways Calculation Of Energy Yields OCN Biogeochemical Systems 8/27/09

2 Earth s History (continental crust) 170 Oldest oceanic crust Ga = billions of years ago

3 The Traditional Approach

4 A Modern Approach Determined from sequencing of ribosomal RNA (as opposed to morphology and physiology) Bacteria Archaea Eucaryotes Crenarchaeota Euryarchaeota Last universal common ancestor Archaea most primitive, live in extreme environments Modified from Chan et al. (1997)

5 Bacteria Archaea Eucaryotes Crenarchaeota Euryarchaeota The lengths of the branches reflect how much the DNA of each lineage has diverged from their common ancestor Conclusions: Most of life's genetic diversity is microbial Multicellular organisms such as fungi, plants and animals evolved from unicellular organisms further down the tree

6 A further complication: endosymbiosis See handout for further information on the five branches of the Tree of Life

7 Metabolic Strategies for Sustaining Life We can classify organisms by function -- how they obtain energy and cell-carbon: 1. Method of energy generation (reactions that convert ADP to ATP): Photosynthesis -- Phototroph e.g.: Oxic photosynthesis: 6 CO H 2 O C 6 H 12 O O 2 + ATP Oxidation/reduction of inorganic compounds -- Lithotroph e.g.: Ammonia oxidation: NH ½ O 2 NO H + + H 2 O + ATP Oxidation of organic compounds -- Organotroph e.g.: Glucose oxidation: C 6 H 12 O O 2 6 CO H 2 O + ATP 2. Carbon source: Carbon dioxide -- Autotroph Organic compounds -- Heterotroph

8 Metabolic Strategies - Examples Photoautotrophs Green plants Cyanobacteria Autolithotrophs Most algae Some purple and green bacteria Methane oxidizing bacteria: CH 4 + O 2 CO 2 + 4H + + 4e - Hydrogen oxidizing bacteria Iron oxidizing bacteria Nitrifying bacteria: NO ½ O 2 NO - 3 Photoheterotrophs Most purple and green bacteria Some algae Some cyanobacteria Heterolithotrophs Sulfide oxidizing bacteria Organoheterotrophs Animals Most bacteria Fungi Protozoa

9 Aerobic and Anaerobic Modes of Organic Matter Respiration (Organoheterotrophy) In general: Reduced organic matter + oxidant CO 2 + reduced oxidant + energy ΔG r Mode of organic matter oxid Oxidant Reduced Oxidant (kj/mole) Aerobic respiration O 2 H 2 O Manganese reduction MnO 2 Mn Nitrate reduction HNO 3 N Iron reduction Fe 2 O 3 Fe FeOOH Fe Sulfate reduction SO 4 2- S Methanogenesis CO 2 CH 4-350

10 Linkage of Oxidations and Reductions Oxidized Reduced CH 2 O generic organic matter These are the most common redox reactions; many others are possible Reduced Oxidized CH 2 O CO 2 Aerobic Respiration O 2 H 2 O H 2 O O 2 NH 4 NO 3 H 2 S SO 4 CO 2 CH 2 O Oxygenic Photosynthesis Chemoautotrophic nitrification CO 2 CH 2 O Chemoautotrophic sulfur oxid CO 2 CH 2 O CH 2 O CO 2 Denitrification NO 3 N 2 CH 2 O CO 2 Anaerobic respiration SO 4 H 2 S In this course we will see that global biogeochemical cycles are driven by transformations of substances between oxidized and reduced forms in different environments

11 Calculation of Energy Yields Consider this general reaction: aa + bb cc + dd The free-energy change of the reaction at standard state can be calculated as: ΔG r = ( cg + dg ) ( ag + bg ) C D A B where G X is the free energy of formation for a product or reactant X at standard state (which can be looked up) Standard state: 1 atm pressure 25 C Concentrations = 1 mole/kg

12 If ΔG r < 0, then the reaction proceeds spontaneously as written (at standard state) If ΔG r > 0, then the reaction proceeds spontaneously in the opposite direction as written (at standard state) If ΔG r = 0, then the reaction is at equilibrium and will not proceeds spontaneously in either direction (at standard state) Thus, ΔG r is a powerful tool to predict if a reaction will occur! However, we are commonly interested in conditions other than standard state...

13 We calculate the in-situ ΔG r using this equation: In-situ ΔG Standard state = ΔG r r + RT ln [ ] c [ ] d C D [ ] a A [ B] b R = ideal gas constant = cal K -1 mol -1 = 8.31 J K -1 mol -1 T = K [ ] = concentration Thus, using the same criteria used on the previous slide, we can predict whether a reaction will occur under real-world conditions! See the Energetics handout for further details

14 The next lecture: Atmospheric deposition, atmospheric models This will begin our investigation of biogeochemical systems on Earth: Atmosphere Terrestrial systems Aquatic systems Lakes Streams and rivers Estuaries Oceans

Lecture 2 Carbon and Energy Transformations

1.018/7.30J Fall 2003 Fundamentals of Ecology Lecture 2 Carbon and Energy Transformations READINGS FOR NEXT LECTURE: Krebs Chapter 25: Ecosystem Metabolism I: Primary Productivity Luria. 1975. Overview