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Anaerobic Respiration
William J Payne, University of Georgia, Athens, Georgia, USA
Anaerobic respiration is growth-supporting utilization by some archaea, bacteria and filamentous fungi of nitrogen oxides, carbon dioxide, sulfate or various metals and organic materials as terminal oxidants in place of oxygen. In nature, the processes exert significant ecological and economic impacts that may be either beneficial or deleterious.

Secondary article
Article Contents
. Introduction . What is Anaerobic Respiration? . How Does Anaerobic Respiration Differ from Aerobic? . Distribution of Anaerobic Respiration among Micoorganisms . Ecological Impact of Anaerobic Respiration

Introduction
Although all sorts of organisms respire, most denitions of respiration describe the act of breathing, the distribution of O2 to the tissues and the exhalation of CO2. Characteristic oxidativereductive, energy-yielding reactions are often noted only as an afterthought, but it is these reactions that actually constitute respiration. This article focuses on the metabolic processes of microorganisms that respire without involving gaseous O2 and grow at the expense of the energy thus conserved.

. Economic Impact of Sulfate Reduction and Denitrification

How Does Anaerobic Respiration Differ from Aerobic?


Energy yield
Transfer of electrons from substrates to a terminal oxidant during anaerobic respiration yields less conservable energy than does transfer to O2. For example, complete aerobic oxidation of glucose releases 2813 kJ mol 2 1, but similar oxidation by nitrate yields only  67% of that amount. Molar energy yields for oxidation by fumarate, sulfate and CO2 are much smaller. Molar biomass yields for microorganisms growing by anaerobic respiration are thus less than those for microbes utilizing aerobic respiration. Both these respiratory types dier from fermentation in two salient aspects: fermenting organisms make use of degradation fragments of their organic substrates as terminal oxidant, thus leaving reduced organic end products (e.g., glucose to pyruvate (the fragment) to lactate (the end product)), and conserve energy by substrate-level, not oxidative, phosphorylation. Growth yields for fermentation are typically smaller than those for aerobic respiratory systems or the higher-potential anaerobic respiratory systems.

What is Anaerobic Respiration?


In various organisms, respiration comprises several sets of linked, enzyme-mediated, electron-transfer pathways whereby reduced substrates are oxidized stepwise, with accompanying reduction of intermediate coenzymes that pass electrons along by stages to acceptance by a terminal oxidant: O2 in aerobic respiration; one or more of an array of elemental, inorganic or organic substances in the alternative, anaerobic respiration. These pathways are coupled at certain electron-transfer points to energyconserving translocation of protons across membranes. That translocation drives production of high-energy phosphates, such as adenosine triphosphate (ATP), that supply energy for cellular biosynthesis and maintenance. Higher eukaryotes oxidize a relatively limited range of reduced substrates, for example a number of sugars, amino acids and lipids; but prokaryotes and lower eukaryotes utilize not only these but a vast array of other materials, including H2, NH3, oligo-organics of natural or xenobiotic origin, hydrocarbons, biopolymers and detergents even heavily halogenated aromatics as well. Anaerobic respiration leading to reduction of higher-electropotential terminal oxidants such as nitrogen oxides (Table 1) makes use of several inducible enzymes that are synthesized in ordinarily aerobic but facultative organisms only in response to limitation or absence of O2. Fewer such enzymes occur in strictly anaerobic sulfate reducers and acetogenic and methanogenic prokaryotes (CO2 reducers).

Electron-transfer enzymes and cofactors utilized (Table 2)


Reduction of nitrate to ammonia as prelude to nitrogen assimilation is a capacity common to plants, fungi, algae and many bacteria in need of nutrient nitrogen and is not 2 anaerobic respiration. Dissimilatory NO3 reduction is a widespread type of anaerobic respiration governed by availability of oxygen and not by nutrient nitrogen supply. It is typically initiated by induced nitrate reductase (NaR), a complex membrane-bound enzyme comprising a cytochrome b, an ironsulfur peptide, and a molybdoprotein as catalytic site. In many microorganisms, a multi (46)-haem cytochrome-c nitrite reductase (NiR) reduces the nitrite
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Table 1 Terminal oxidants utilized by microbes for anaerobic respiration (adapted from Madigan et al., 1997) Couples Inorganic aceptors 2 32 AsO3 4 /AsO3 2 ClO3 /Cl 2 CO2/CH4 CO2/acetate Fe3 1 /Fe2 1 Mn4 1 /Mn2 1 2 2 /NO2 NO3 2 NO2 /NO NO/N2O N2O/N2 2 22 SeO2 4 /SeO3 0 2 S /HS 2 2 SO2 4 /HSO3 2 (APS/HSO3 1 AMP) 2 2 SO2 4 /HS 2 HSO3 /HS 2 2 2 2 1 HSO3 S2O2 3 /HS Organic acceptors Dimethyl sulfoxide/dimethyl sulde Fumarate/succinate Glycine/acetate 1 ammonia Halogenated phenols Humic substances Trimethylamine N-oxide/trimethylamine
a

E0 (V)
a

1 0.139 + 1.03 2 0.24 2 0.29 1 0.771 1 0.798 1 0.43 1 0.36 1 1.18 1 1.36 1 0.475 2 0.27 2 0.52 ( 2 0.06) 2 0.22 2 0.11 2 0.40

1 1 2 1

0.16 0.03 0.01

0.13

E0 (V) for O2/H2 is 1 0.815, less than the values for NO/N2O and N2O/N2. Few coupling sites exist in chains leading to reduction of N2O, however, and molar growth yields for this terminal oxidant are small. Use of NO as sole terminal oxidant for growth is questionable, occurring if anywhere in Thiobacillus denitricans. Cytochrome c3 (ox/red) has a redox potential of 2 0.29 V and 2 is activated by ATP-sulfurylase. reduces the anion only after SO2 4

produced to ammonia. This enzyme can also reduce nitric oxide to ammonia, if the NiR is completely reduced, or to N2O, if the NiR is only partially reduced. Denitrication (that type of anaerobic respiration whereby some microorganisms can reduce nitrate seriatim to nitrite, nitric oxide, nitrous oxide and N2) also makes use of an induced NaR that branches electron transfer at the quinone of the 2 2 to NO2 . usual electron transport chain to reduce NO3 2 For reduction of NO2 several prokaryotes and a few lamentous fungi then move electrons further along from the quinone to cytochrome bc1 and branch electron transfer after the cytochrome bc1 complex. At this branch point, some denitriers produce a copper protein NiR that reduces nitrite. Other prokaryotes make use of a tetrahaem cytochrome NiR containing two c and two d1 haems. None produces both types of NiR. Type II Cu atoms and the d1 haem are the catalytic sites for these induced NiR. Nitric oxide, NO, the product of both
2

denitrifying NiR types, serves dually as reducible intermediate and as activator of the gene nnrR, regulating both NiR and nitric oxide reductase (NoR) production and function (Tosques et al., 1996). The extremely reactive nature of NO requires that its production and reduction rates be closely regulated to prevent nonspecic side reactions. Membrane-bound NoR is a cytochrome bc complex in prokaryotes but a cytochrome P450 in denitrifying lamentous fungi. Some denitriers stop the process with N2O production; most complete the N2 cycle by use of an 8-Cu protein N2O reductase (NNoR), which is structurally akin to cytochrome c oxidase. NNoR contain copper of two sorts: mixed-valence CuA as electron carrier and a binuclear copper centre CuZ as catalytic site. NNoR can be nondestructively inhibited by acetylene. This property facilitates environmental assay by gas chromatographic measurement of accumulating N2O. In many denitriers, c-type cytochromes are found to receive electrons from the bc1 complex and transfer them to the NiR, NoR and NNoR, although blue-copper proteins are donors for the Cu-NiR instead of such cytochromes in several denitriers (Zumft, 1997). With a redox couple greater than 1 0.35 V for each reduction step, denitriers need not carry out every step in the process to achieve net conservation of energy. With an E0 (V) of 1 0.03, fumarate can receive electrons from reduced nicotinamide and avin coenzymes of the electron transport chain. Anaerobically respiratory fumarate-reducing bacteria thus utilize membrane-bound fumarate reductase, a avinadenine dinucleotide (FAD) enzyme that also contains three ironsulfur clusters, with electrons supplied by formate dehydrogenase. Other bacteria couple reduction of fumarate to oxidation of reduced nicotinamideadenine dinucleotide. Succinate is released and energy for ATP production is conserved. Unlike nitrate, sulfate must be activated before biological reduction to sulte takes place. This is necessary as the redox potential of the sulfate/bisulte couple is 2 0.516 V; that of the biologically powerful reductant 2H 1 /H2 is 2 0.414 V. Sulfate must therefore be rendered reducible by expenditure of energy-rich ATP. Assimilatory reduction of sulfate that supplies sulfur for amino acid biosynthesis in a variety of organisms requires expenditure of two ATP to generate adenosine 5-phosphosulfate, the substrate for reduction. In contrast, activation of sulfate by ATPsulfurylase in preparation for dissimilatory anaerobic respiration in sulfate-reducing bacteria (SRB) consumes one ATP to produce adenosine 5-phosphosulfate (APS), the terminal oxidant for the FAD, 8-Fe enzyme, APSreductase. The pyrophosphate released from ATP during this reaction must be hydrolysed to provide thermodynamic stability. Molecular hydrogen arising from fermentation in the environment or from SRB catabolism of a wide variety of organic compounds, reduces the periplasmic enzyme, hydrogenase, which transfers electrons to the (E0 electronegative tetrahaem cytochrome c3

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Table 2 Some aspects of better-known anaerobic respiratory systems Respiratory process Nitrate reduction/ ammonification Electron donors Electron acceptors

Products NO2

Typical organisms Escherichia coli

Various organic and NO3 inorganic compounds

Nitrate reductase (molybdoprotein)

NO2

Nitrite reductasea (multi-(4-6) haem c cytochrome)

NH3

Wolinella succinogenes

Denitrification

Various organic and inorganic compounds and elements such as H2

NO3

Nitrate reductase (molybdoprotein)

NO2

Ralstonia eutropha

NO2

Nitrite reductase (cytochrome cd l) or (6-Cu trimeric protein) c

NO b

Thiobacillus denitrificans or Bacillus halodenitrificans

2NO

Nitric oxide reductase (cytochrome bc)

N2O

Pseudomonas stutzeri

N2O

Nitrous oxide reductase (8-Cu protein)

N2

Paracoccus denitrificans

Fumarate reduction

Formate

Fumarate
Fumarate reductase (FAD-linked protein)

Succinate

Proteus rettgeri

Sulfate reduction

Lactate, glycerol, etha- SO42 nol, aliphatic and aromatic hydrocarbons HSO3

APS reductase d

HSO3

Sulfite reductase

H2S CH4

Desulfovibrio gigas Methanosarcina barkeri

Methanogenesis

H2, formate, acetate

CO2

(coenzyme-bound C 1 intermediates)

a b

Can also reduce NO. Exerts regulatory influence on production and functioning of nitrite and nitric oxide reductases as well. c This type of nitrite reductase is also produced by the denitrifying filamentous fungi and actinomycetes. d Initial reduction requires input of energy for activation. The processes supply net energy for conservation by reduction of sulfite and subsequently formed intermediates.

(V) 5 2 0.29) in the membrane while extruding two protons. Reduced c3 supplies electrons via carrier proteins 2 1 AMP, for reduction of APS to yield sulte (APS/HSO3 E0(V) 5 2 0.06). The (bi)sulte produced by APS reduction is then reduced to sulde with net ATP gain. The

reduction is carried out by one of four bisulte reductases: desulfoviridin, desulfofuscidin, P590 or desulforubidin in various SRB. Although SRB grow by anaerobic respiration only, some can couple O2 reduction to phosphorylation as an apparent survival mechanism. SRB that oxidize
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acetate use a modied citric acid cycle and produce one ATP from functioning of the novel enzyme citrate lyase or the key enzyme CO dehydrogenase (CODH). SRB also produce various redox components such as mono-, octaand hexadecahaem cytochromes c, rubredoxin, avodoxin, rubrerythrin, ferredoxin and cytochrome b, several with yet uncertain function (Chen et al., 1995). Methanogenic archaea are the strictest of all anaerobes. To generate ATP, they reduce CO2 at the expense of H2, utilizing unique carriers and reductases. CO2 is rst attached to methanofuran (MF) and reduced to the formyl state at the expense of an endergonic reaction dependent upon reverse electron transport. The formyl group is transferred to tetrahydromethanopterin (MP), and reduces seriatim to methenyl, methylene and methyl states, the latter two products at the expense of two H2 that reduce 2 moles of carrier coenzyme F420. The MP donates the methyl group to 2-mercaptoethanesulfonic acid (S-CoM) supplying the energy for the reverse electron transport. The methylreductase system (comprising 7-mercaptoheptanoylthreonine phosphate (HTP-S), which has been reduced by H2 while coupled with S-CoM, and coenzyme F430, a nickel tetrapyrrole) then provides the energyconserving step by reducing the methyl to free CH4 and reconjoining HTP-S-S-CoM. At the expense of four H2, the total free energy change is 2 130.4 kJ mol 2 1 CH4. Methanogens dismutating acetate rst form acetyl-coenzyme A, then oxidize the carbonyl group to CO2, and use the electrons gained to reduce the methyl group to CH4. AcetylCoA interacts with CODH, which retains the carbonyl and transfers the methyl group to a corrinoid carrier that delivers it to CoM, where it is reduced to CH4 (the energy-conserving step) by electrons generated by oxidation of CO linked to CODH, liberating CO2. The total free energy change is 2 36 kJ mol 2 1. Methanol can enter the pathway of methanogenesis by attaching to the corrinoid, undergoing reduction to the methyl state, and transfering to CoM, where energy-conserving reduction to CH4 occurs; the total free energy change is 2 106 kJ mol 2 1. The reaction involves 4 moles methanol and yields 3 moles of CH4 and 1 mole of CO2 (Madigan et al., 1997).

terminal oxidants for several prokaryotes, presumably because of their reducible quinone moieties. The analogue 2,6-anthraquinone disulfonate was recently found to substitute for humic material in laboratory experiments. Microbial reduction of widely available Fe(iii ), with concomitant degradation of much organic matter, yields soluble Fe(ii ) and accumulation of the minerals magnitite and siderite. Similary, reduction of Mn(iv ) generates manganese minerals and supports degradation of organic substances in aquifers and sediments. Reduction of selenate to selenite and selenium supports growth of several organisms; others utilize U(vi) and Tc(vii ), reducing them to insoluble U(iv ) and several Tc species, respectively. Reduction of chlorate to chlorite ordinarily kills microorganisms, but bacteria capable of growth at the expense of this reaction have been isolated. Elemental sulfur serves exclusively as terminal oxidant for growth of certain archaea and may also be used as terminal oxidants by several sulfate-reducing bacteria (Lovley, 1993).

Distribution of Anaerobic Respiration among Micoorganisms


Ability to grow at the expense of anaerobic respiration appears widely scattered in the microbial world. Capacity for induced nitrate-reducing anaerobic respiration that stops at nitrite or proceeds to ammonia production may be the most frequently encountered type of anaerobic respiration in the biosphere. The property is exhibited by prokaryotes as diverse as Escherichia coli and other enterics, Mycobacterium tuberculosis, Staphylococcus intermedius, Pasteurella species, Clostridium, Wolinella and Desulfovibrio species, as well as many others. Several archaea carry out denitrifying anaerobic respiration in harsh environments. Included are extreme halophiles, Haloarcula and Haloferax, and hyperthermophiles, Pyrococcus and Ferroglobus species. The chemolithotrophic organism Ferroglobus is perhaps unique as a strictly, as opposed to facultatively, anaerobic denitrier. Many proteobacteria exhibit denitrication. In the a subclass, organisms as diverse as phototrophic Rhodobacter and budding Hyphomicrobium species, diazotrophic Azospirillum and Rhizobium strains, and both facultatively chemolithotrophic Paracoccus and constitutive Nitrobacter species denitrify. Denitrifying members of the b subclass include broadly dispersed Alcaligenes and Nitrosomonas species, Thiobacillus strains and several Neisseria isolates such as the notable pathogens N. gonorrhoeae and N. meningitidis. g-Subclass denitriers are the most frequently isolated and include several Pseudomonas species. A few are psychrophiles. Strains of the complex Beggiatoa, Thermothrix and Flexibacter species are denitriers. Gram-positive denitriers are found among the actinomycetes and in the genus Bacillus. These include

Array of additional terminal electron acceptors utilized


In addition to the nitrogen oxides, fumarate, sulfate and CO2, several other organic compounds, metals and oxyanions can also serve as terminal oxidants for anaerobic respiration (Table 1). Natural organic products in sh, dimethyl sulfoxide and trimethylamine N-oxide, can also act as terminal oxidants for many prokaryotes, making use of a molybdoprotein reductase and leading to release of odoriferous products. Halogenated phenols undergo reductive dehalogenation while serving as terminal oxidants for some bacteria. Humic substances serve as
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halophilic B. halodenitricans and thermophilic B. stearothermophilus. In all, as many as 50 genera of usually aerobic, but facultative, prokaryotes include denitrifying species. Until recently, denitrication was thought to be restricted to the prokaryotes, but a few lamentous fungi in the genera Fusarium and Cylindrocarpon were also shown to denitrify utilizing mitochondrial enzymes and stopping the process at N2O (Zumft, 1997). Facultative fumarate respiration occurs broadly in nature, for example in Halobacterium and Haloferax strains among archaea and in E. coli, Fibrobacter and Wolinella among several additional eubacteria (Meinhardt and Glass, 1994). Bacteria reducing humic materials t into the family Geobacteraceae. All reduce Fe(iii ) as well. bSubclass proteobacterial isolates reducing chlorate, perchlorate and nitrate constitute a new genus, Dechlorimonas. Morphologically diverse prokaryotes that generate sulde from sulfate reduction in anoxic waters are assigned to the archaean genus Archaeoglobus or to the eubacterial genera such as Desulfovibrio, Desulfobacter, Desulfosarcina, Thermodesulfobacterium and Desulfotomaculum. This last-mentioned organism is a Gram-positive endosporeformer. Members of the genus Desulfomonas are found in human intestines. Archaean thermophiles in the genera Acidianus, Pyrobaculum and Pyrodictium utilize sulfur as terminal oxidant, as do thermophilic eubacteria in the genera Desulfuromonas, Desulfurella and Thermotoga. Several sulfur-reducing members of the order Thermotogales reduce thiosulfate as well, but not sulfate. A few Desulfovibrio and Desulfobacter species are diazotrophic (Stackebrandt et al., 1995). Methanogens are grouped in ve orders within Archaea. The mainly rod-shaped species are assigned to the Methanobacteriales, which includes thermophiles and the extremely thermophilic Methanothermus strains among the many mesophiles. Marine coccoid methanogens comprise the order Methanococcales, which also contains thermophiles (e.g. Methanothermococcus thermolithotrophicus). Requirement for acetate as carbon source characterizes the Methanomicrobiales, whose members vary morphologically and include a sheathed species, Methanospirillum hungateii. The aggregated cocci and macrocysts such as Methanosarcina species that catabolize various methyl-group compounds such as methanol, methyl amines and methyl sulde comprise the order Methanosarcinales. A single species, Methanopyrus kandleri, constitues the order Methanopyrales, an organism restricted to catabolizing H2 and reducing only CO2 while growing at or about boiling water temperature (Boone et al., 1993). Rather than methanogenic, two dierent types of anaerobic bacteria, Clostridium aceticum and Acetobacterium woodii, are acetogenic. They reduce CO2 while oxidizing H2 and yield acetate rather than CH4 as the nal product.

Versatility characterizes microorganisms carrying out anaerobic respiration with several terminal oxidants. The obligately anaerobic d-proteobacterium Geobacter metallireducens utilizes Fe(iii ), Mn(iv), U(vi ) or nitrate. Perhaps the most versatile of all is the facultatively anaerobic dproteobacterium Shewanella putrefaciens, which can grow at the expense of reduction of Fe(iii ), Mn(iv ), nitrate, nitrite, thiosulfate, sulfur, technetium, trimethylamine Noxide and fumarate. Somewhat less versatile are anaerobic Gram-positive Fe(iii )-reducers designated Thermoterrabacterium and sulfur-reducing and chlorophenol-reducing Desultobacterium species, as well as an unidentied bacterium that respires not only with several of these terminal oxidants but also with arsenate (Lovley, 1993).

Ecological Impact of Anaerobic Respiration


Setting aside questions whether anaerobic respiration or aerobic respiration developed rst as life evolved, and many hold anaerobic respiration as primal, the capacity for anaerobic respiration permits microorganisms to occupy and thrive in a remarkable variety of oxygen-poor or anoxic niches, from ice to boiling water, oral cavities to groundwater, sewage to silos. For convenience, the contributions of anaerobic respiration to complexity of life on earth can be examined under three headings.

Biodiversity
Demonstration of the expansive diversity of anaerobic respiration-utilizing microorganisms greatly enlarges our perception of the range of capability of DNA to establish and maintain life on earth, and perhaps beyond.

Recycling
Recognition of the existence of anaerobic respiration in all its forms increases awareness of the number and variety of mechanisms functional in recycling of xed elements needed for biosynthesis by succeeding generations, notably carbon, nitrogen and sulfur. Mineralization (carbon cycling) of both complex and simple organic matter by anaerobic respiration in sediments, soil, aquifers and rocks provides energy and nutrients that sustain a massive subsurface microbial biomass thought by some to outweigh in aggregate all above-ground biota. Extensive anaerobic respiration-driven carbon cycling also takes place in bodies of water and many other above-ground environments, including animal rumina and intestines. Farmers see nitrogen cycling by denitriers as a defect that lessens soil fertility, but removal of wastewater nitrogen after either ammonication and nitrication or agricultural nitrate runo aids in prevention of widespread
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eutrophication in lakes and streams. Release of CO2, CH4 and nitrogen oxides during or as a consequence of anaerobic respiration contributes to normal, equilibriumpromoting erosion of the stratospheric ozone layer; but practices that increase the quantities of these gases liberated may contribute to global warming and enhanced penetration of the earths atmosphere by UV irradiation. Sulfate reduction serves for bioremediation of acid mine waters and enhances carbon cycling in microbial mats and sediments but adds to corrosion and fouling in submerged equipment that plagues such enterprises as oil recovery and distribution.

Linking energy conservation to metal metabolism


Use of metals as terminal oxidants for anaerobic respiration changes their physical state and encourages deposition of some (e.g., U(vi ) to U(iv ); selenate to selenite and selenium; Tc(vii ) to Tc(iv )) and solubilization of others (e.g., Fe(iii ) to Fe(ii) and Mn(iv ) to Mn(ii), reduced products that can react with sulde to form precipitable or insoluble compounds).

that represents an economic burden. Fixed nitrogen is usually the limiting nutrient for crop and plant growth in nature, and replacement of lost nitrogen by microbial xation, mining of nitrate or manufacture of fertilizer is likely to be expensive. Estimates of costs of nitrogen losses to denitrication as great as 30% from applied nitrate fertilizer before crop plants can assimilate the nutrient are consistent with that notion, particularly in view of the expense of mining, packaging, shipping, distributing and applying nitrate. On the positive side, much plant residue is degraded (carbon cycling) and a great deal of soil conditioning takes place during denitrication. Exploitation of denitrication for simultaneous removal of organic accumulation and heavy nitrogen loading in feed-lot runo water, in sewage undergoing treatment, and in many industrial waste waters subjected to remediation may further oset negative economic eects of the phenomenon. Whether it is viewed as asset or defect, no reasonable alternatives to denitrication exist for the daily handling at acceptable cost of the huge volumes of such wastes as our civilization generates (Payne, 1981).

References
Boone DR, Whitman WB and Rouviere P (1993) Diversity and taxonomy of methanogens. In: Ferry JG (ed.) Methanogenesis Ecology, Physiology, Biochemistry and Genetics, pp. 3580. New York: Chapman and Hall. Chen L, Liu M-Y and LeGall J (1995) Characterization of electron transfer proteins. In: Barton LL (ed.) Sulfate-Reducing Bacteria, pp. 113149. New York: Plenum Press. Hamilton WA and Lee W (1995) Biocorrosion. In: Barton LL (ed.) Sulfate-Reducing Bacteria, pp. 243264. New York: Plenum Press. Lovley DR (1993) Dissimilatory metal reduction. Annual Review of Microbiology 47: 263290. Madigan MT, Martinko JM and Parker J (1997) Brock Biology of Microorganisms, 8th edn. Upper Saddle River, NJ: Prentice Hall. Meinhardt SW and Glass TL (1994) NADH-linked fumarate reductase and NADH dehydrogenase activities in Fibrobacter succinogenes. Current Microbiology 28: 247251. Payne WJ (1981) Denitrication. New York: Wiley-Interscience. Stackebrandt E, Stahl DA and Devereux R (1995) Taxonomic relationships. In: Barton LL (ed.) Sulfate-Reducing Bacteria, pp. 4987. New York: Plenum Press. Tosques IE, Shi J and Shapleigh JP (1996) Cloning and characterization of nnrR, whose product is required for expression of proteins involved in nitric oxide metabolism in Rhodobacter sphaeroides 2.4.3. Journal of Bacteriology 178: 49584964. Zumft WG (1997) Cell biology and molecular basis of denitrication. Microbiology and Molecular Biology Reviews 61: 533616.

Economic Impact of Sulfate Reduction and Denitrification


The most signicant economic impact of anaerobic respiration by SRB is attributable to corrosion of metals by suldes (usually the pitting of surfaces), for example in oshore oil platform structures and equipment (Hamilton and Lee, 1995). The SRB are also accountable for souring of recovered oil and gas and for plugging geological accesses to the petroleum. The eects are particularly damaging when bulky, microbially generated lms enmeshing SRB and other microorganisms cover structures and coat the interiors of pipes. After suldes are formed, exposure to oxygen greatly intensies corrosive eects as pyrite and elemental sulfur are produced. Very expensive and unrelenting eorts at controlling ow-impeding biolm formation include use of plastic pipe liners, dehydration of owing gas, cleansing with biocides such as quaternary ammonium salts, heat shocks and bioelectric eects. Understanding the ecological interrelationships among the SRB and other microbes in biolms and sorting out their signalling processes are critical to prospects for controlling the destructive eects of sulde and biolm formation and their direct and indirect consequences. Unlike sulfate reduction, in which the product of anaerobic respiration exerts deleterious eects, denitrication releases a bland end product (N2) into an atmosphere already comprising  80% N2. It is the loss of nitrate, the choice plant nutrient, to denitrifying anaerobic respiration
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Further Reading
Berks BC, Ferguson SJ, Moir JWB and Richardson DJ (1995) Review. Enzymes and associated electron transport systems that catalyze the respiratory reduction of nitrogen oxides and oxyanions. Biochimica et Biophysica Acta 1232: 97173. Chanton JP and Dacey JWH (1991) Eects of vegetation on methane ux, reservoirs, and carbon isotopic composition. In: Sharkey TD,

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Holland EA and Mooney HA (eds) Trace Gas Emissions by Plants, pp. 6592. San Diego: Academic Press. Hansen TA (1994) Metabolism of sulfate-reducing prokaryotes. Antonie van Leeuwenhoek 66: 165184.

Nealson KH and Saarini DA (1994) Iron and manganese in anaerobic respiration: environmental signicance, physiology, and regulation. Annual Review of Microbiology 48: 311343. Whitman WB and Rogers JE (1991) Microbial Production and Consumption of Greenhouse Gases. Washington, DC: American Society for Microbiology.

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