24.1 The Carbon Cycle Carbon is cycled through all of Earth’s major carbon reservoirs Includes atmosphere, land, oceans, sediments, rocks, and biomass.

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24.1 The Carbon Cycle Carbon is cycled through all of Earth’s major carbon reservoirs Includes atmosphere, land, oceans, sediments, rocks, and biomass Reservoir size and turnover time are important parameters in understanding the cycling of elements © 2012 Pearson Education, Inc.

Figure 24.1 CO2 Figure 24.1 The carbon cycle. Human activities Respiration Land plants Animals and microorganisms Aquatic plants and phyto- plankton CO2 Aquatic animals Biological pump Fossil fuels Humus CO2 Death and mineralization Figure 24.1 The carbon cycle. Soil formation Earth’s crust Rock formation © 2012 Pearson Education, Inc.

24.1 The Carbon Cycle CO2 in the atmosphere is the most rapidly transferred carbon reservoir CO2 is fixed by photosynthetic land plants and marine microbes CO2 is returned to the atmosphere by respiration as well as anthropogenic activities Microbial decomposition is the largest source of CO2 released to the atmosphere © 2012 Pearson Education, Inc.

24.1 The Carbon Cycle Carbon and oxygen cycles are linked Phototrophic organisms are the foundation of the carbon cycle Oxygenic phototrophic organisms can be divided into two groups: plants and microorganisms Plants dominant organisms of terrestrial environments Microorganisms dominate aquatic environments © 2012 Pearson Education, Inc.

24.1 The Carbon Cycle Photosynthesis and respiration are part of redox cycle (Figure 24.2) Photosynthesis CO2 + H2O (CH2O) + O2 Respiration (CH2O) + O2 CO2 + H2O The two major end products of decomposition are CH4 and CO2 © 2012 Pearson Education, Inc.

(CH2O)n CO2 (CH2O)n Oxygenic photosynthesis Respiration Figure 24.2 (CH2O)n Organic matter Oxygenic photosynthesis Respiration Chemolithotrophy Methanotrophy Oxic CO2 Anoxic Methanogenesis Figure 24.2 Redox cycle for carbon. Acetogenesis Anaerobic respiration and fermentation Syntroph assisted Anoxygenic photosynthesis Organic matter (CH2O)n © 2012 Pearson Education, Inc.

24.2 Syntrophy and Methanogenesis Methanogenesis is central to carbon cycling in anoxic environments Most methanogens reduce CO2 to CH4 with H2 as an electron donor; some can reduce other substrates to CH4 (e.g., acetate) Methanogens team up with partners (syntrophs) that supply them with necessary substrates © 2012 Pearson Education, Inc.

24.3 The Nitrogen Cycle Nitrogen A key constituent of cells Exists in a number of oxidation states © 2012 Pearson Education, Inc.

Figure 24.7 Figure 24.7 Redox cycle for nitrogen. Nitrification groups of protein Assimilation Assimilation Nitrogen fixation Ammonification DRNA Oxic Anoxic groups of protein Assimilation Ammonification Nitrogen fixation Anammox Figure 24.7 Redox cycle for nitrogen. Denitrification © 2012 Pearson Education, Inc.

24.3 The Nitrogen Cycle N2 is the most stable form of nitrogen and is a major reservoir Only a few prokaryotes have the ability to use N2 as a cellular nitrogen source (nitrogen fixation) Denitrification is the reduction of nitrate to gaseous nitrogen products and is the primary mechanism by which N2 is produced biologically Ammonia produced by nitrogen fixation or ammonification can be assimilated into organic matter or oxidized to nitrate © 2012 Pearson Education, Inc.

II. Biodegradation and Bioremediation 24.7 Microbial Leaching 24.8 Mercury Transformations 24.9 Petroleum Biodegradation and Bioremediation 24.10 Xenobiotics Biodegradation and Bioremediation © 2012 Pearson Education, Inc.

24.7 Microbial Leaching Bioremediation Refers to the cleanup of oil, toxic chemicals, or other pollutants from the environment by microorganisms Often a cost-effective and practical method for pollutant cleanup © 2012 Pearson Education, Inc.

24.7 Microbial Leaching Microbial leaching The removal of valuable metals, such as copper, from sulfide ores by microbial activities (Figure 24.15) Particularly useful for copper ores © 2012 Pearson Education, Inc.

24.7 Microbial Leaching In microbial leaching, low-grade ore is dumped in a large pile (the leach dump) and sulfuric acid is added (Figure 24.16) The liquid emerging from the bottom of the pile is enriched in dissolved metals and is transported to a precipitation plant Bacterial oxidation of Fe2+ is critical in microbial leaching as Fe3+ itself can oxidize metals in the ores © 2012 Pearson Education, Inc.

Figure 24.16 Sprinkling of acidic solution on CuS Low-grade copper ore (CuS) Copper ore can be oxidized by oxygen- dependent (1) and oxygen-independent (2) reactions, solubilizing the copper: 1. CuS  2 O2  Cu2  SO42 2. CuS  8 Fe3  4 H2O Cu2  8 Fe2  SO42  8 H Soluble Cu2 Acidic solution pumped back to top of leach dump Cu2 H2SO4 addition Figure 24.16 Arrangement of a leaching pile and reactions in the microbial leaching of copper sulfide minerals to yield metallic copper. Recovery of copper metal (Cu0) Fe0  Cu2  Cu0  Fe2 (Fe0 from scrap steel) Acidic Fe2- rich solution Precipitation pond Leptospirillum ferrooxidans Acidithiobacillus ferrooxidans Oxidation pond Copper metal (Cu0) © 2012 Pearson Education, Inc.

24.7 Microbial Leaching Microorganisms are also used in the leaching of uranium and gold ores (Figure 24.17) © 2012 Pearson Education, Inc.

24.9 Petroleum Biodegradation and Bioremediation Prokaryotes have been used in bioremediation of several major crude oil spills (Figure 24.20) © 2012 Pearson Education, Inc.

Figure 24.20 Figure 24.20 Environmental consequences of large oil spills and the effect of bioremediation. © 2012 Pearson Education, Inc.

24.9 Petroleum Biodegradation and Bioremediation Diverse bacteria, fungi, and some cyanobacteria and green algae can oxidize petroleum products aerobically Oil-oxidizing activity is best if temperature and inorganic nutrient concentrations are optimal Hydrocarbon-degrading bacteria attach to oil droplets and decompose the oil and disperse the slick (Figure 24.21) © 2012 Pearson Education, Inc.

Bacteria Oil droplets Figure 24.21 Figure 24.21 Hydrocarbon-oxidizing bacteria in association with oil droplets. © 2012 Pearson Education, Inc.

24.9 Petroleum Biodegradation and Bioremediation Gasoline and crude oil storage tanks are potential habitats for hydrocarbon-oxidizing microbes (Figure 24.22) If sufficient sulfate is present, sulfate-reducing bacteria can grow and consume hydrocarbons © 2012 Pearson Education, Inc.

Figure 24.22 Figure 24.22 Bulk petroleum storage tanks. © 2012 Pearson Education, Inc.

24.9 Petroleum Biodegradation and Bioremediation Some microorganisms can produce petroleum Particularly certain green algae © 2012 Pearson Education, Inc.

24.10 Xenobiotics Biodegradation and Bioremediation Xenobiotic compound Synthetic chemicals that are not naturally occurring Examples: pesticides, polychlorinated biphenyls, munitions, dyes, and chlorinated solvents (Figure 24.23) Many degrade extremely slowly © 2012 Pearson Education, Inc.

DDT, dichlorodiphenyltrichloroethane (an organochlorine) Figure 24.23 DDT, dichlorodiphenyltrichloroethane (an organochlorine) Malathion, mercaptosuccinic acid diethyl ester (an organophosphate) Site of additional Cl for 2,4,5,-T 2,4-D, 2,4-dichlorophenoxy- acetic acid Atrazine, 2-chloro-4-ethylamino- 6-isopropylaminotriazine Figure 24.23 Examples of xenobiotic compounds. Monuron, 3-(4-chlorophenyl)- 1,1-dimethylurea (a substituted urea) Chlorinated biphenyl (PCB), shown is 2,3,4,2,4,5- hexachlorobiphenyl Trichloroethylene © 2012 Pearson Education, Inc.

24.10 Xenobiotics Biodegradation and Bioremediation Pesticides Common components of toxic wastes Include herbicides, insecticides, and fungicides Represent a wide variety of chemicals Some can be used as carbon sources by microorganisms Some can be used as electron donors © 2012 Pearson Education, Inc.

24.9 Xenobiotics Biodegradation and Bioremediation Some xenobiotics can be degraded partially or completely if another organic material is present as a primary energy source (cometabolism) © 2012 Pearson Education, Inc.

24.10 Xenobiotics Biodegradation and Bioremediation Chlorinated xenobiotics can be degraded anaerobically (reductive dechlorination) or aerobically (aerobic dechlorination) Reductive dechlorination is usually a more important process as anoxic conditions develop quickly in polluted environments © 2012 Pearson Education, Inc.

24.10 Xenobiotics Biodegradation and Bioremediation Plastics of various types are xenobiotics that are not readily degraded by microorganisms (Figure 24.25) The recalcitrance of plastics has fueled research efforts into a biodegradable alternative (biopolymers) © 2012 Pearson Education, Inc.

Polyvinyl chloride (PVC) Figure 24.25 Polyethylene Polypropylene Polyvinyl chloride (PVC) Polystyrene Polyurethane Teflon PHV PHB Figure 24.25 Synthetic and microbial plastics. © 2012 Pearson Education, Inc.