Fatty acids and alcohols are key intermediates in the methanogenic degradation of organic matter, e.g., in anaerobic sewage sludge digestors or freshwater lake sediments. They are produced by classical fermenting bacteria for disposal of electrons derived in simultaneous substrate oxidations. Methanogenic bacteria can degrade primarily only one-carbon compounds. Therefore, acetate, propionate, ethanol, and their higher homologs have to be fermented further to one-carbon compounds. These fermentations are called secondary or syntrophic fermentations. They are endergonic processes under standard conditions and depend on intimate coupling with methanogenesis. The energetic situation of the prokaryotes cooperating in these processes is problematic: the free energy available in the reactions for total conversion of substrate to methane attributes to each partner amounts of energy in the range of the minimum biochemically convertible energy, i.e., 20 to 25 kJ per mol per reaction. This amount corresponds to one-third of an ATP unit and is equivalent to the energy required for a monovalent ion to cross the charged cytoplasmic membrane. Recent studies have revealed that syntrophically fermenting bacteria synthesize ATP by substrate-level phosphorylation and reinvest part of the ATP-bound energy into reversed electron transport processes, to release the electrons at a redox level accessible by the partner bacteria and to balance their energy budget. These findings allow us to understand the energy economy of these bacteria on the basis of concepts derived from the bioenergetics of other microorganisms.

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Energetics of syntrophic cooperation in methanogenic degradation.

Among different conversion processes for biomass, biological anaerobic digestion is one of the most economic ways to produce biogas from various biomass substrates In addition to hydrolysis of polymeric substances, the activity and performance of the methanogenic bacteria is of paramount importance during methanogenesis The aim of this paper is primarily to review the recent literature about the occurrence of both acetotrophic and hydrogenotrophic methanogens during anaerobic conversion of particulate biomass to methane (not wastewater treatment), while this review does not cover the activity of the acetate oxidizing bacteria Both acetotrophic and hydrogenotrophic methanogens are essential for the last step of methanogenesis, but the reports about their roles during this phase of the process are very limited Despite, some conclusions can still be drawn At low concentrations of acetate, normally filamentous Methanosaeta species dominate, eg, often observed in sewage sludge Apparently, high concentrations of toxic ionic agents, like ammonia, hydrogen sulfide (H2S) and volatile fatty acids (VFA), inhibit preferably Methanosaetaceae and especially allow the growth of Methanosarcina species consisting of irregular cell clumps, eg, in cattle manure Thermophilic conditions can favour rod like or coccoid hydrogenotrophic methanogens Thermophilic Methanosarcina species were also observed, but not thermophilic Methanosaetae Other environmental factors could favour hydrogentrophic bacteria, eg, short or low retention times in a biomass reactor However, no general rules regarding process parameters could be derivated at the moment, which favours hydrogenotrophic methanogens Presumably, it depends only on the hydrogen concentration, which is generally not mentioned in the literature

The roles of acetotrophic and hydrogenotrophic methanogens during anaerobic conversion of biomass to methane: a review

Ammonia inhibition in anaerobic digestion: A review

http://hoffman.cm.utexas.edu/courses/wood_lungdahl_1.pdf

Acetogenesis and the Wood-Ljungdahl Pathway of CO2 Fixation

Biological methanogenesis plays a major role in the carbon cycle on Earth. Methanogenesis is the terminal step in carbon flow in many anaerobic habitats, including marine and freshwater sediments, marshes and swamps, flooded soils, bogs, geothermal habitats, and animal gastrointestinal tracts. CH4 escaping from anaerobic habitats can serve as a carbon and energy source for aerobic methanotrophic bacteria, and can escape to the atmosphere, where it is a major participant in atmospheric chemical reactions and is an important greenhouse gas.

Physiological Ecology of Methanogens

Methanogenesis from acetate by a rod-shaped enrichment culture grown at 60° C was found to require the presence of two organisms rather than a single aceticlastic methanogen. A thermophilic Methanobacterium which grew on H2/CO2 or formate was isolated from the enrichment. Lawns of this methanogen were used to co-isolate an “acetate oxidizer” in roll tubes containing acetate agar. The rod-shaped acetate oxidizer was morphologically distinct from the methanogen and did not show F420 autofluorescence. The coculture completely degraded 40 μmol/ml acetate, and produced nearly equal quantities of methane, and methanogenesis was coupled with growth. The doubling time for the coculture at 60°C was 30–40 h and the yield was 2.7±0.3 g dry wt/mol CH4. Studies with 14C-labelled substrates showed that the methyl group and the carboxyl group of acetate were both converted primarily to CO2 by the coculture and that CO2 was concurrently reduced to CH4. During growth, there was significant isotopic exchange between CO2 and acetate, especially with thecarboxyl position of acetate. These results support a mechanism for methanogenesis from acetate by the coculture in which acetate was oxidized to CO2 and H2 by one organism, while H2 was subsequently used by a second organism to reduce CO2 to CH4. Since the H2 partial pressure must be maintained below 10-4 atm by the methanogen for acetate oxidation to be thermodynamically feasible, this is an example of obligate interspecies hydrogen transfer. This mechanism was originally proposed for a single organism by Barker in 1936.

Markus Koch

Papers

Non-aceticlastic methanogenesis from acetate: acetate oxidation by a thermophilic syntrophic coculture