Anaerobic ammonium oxidation pathway

The biogeochemical cycling of nitrogen is very much controlled by redox reactions. This perspective is presented in Figure 24.3 for the redox reactions that take place in the water column and sediments. The major pathways of reduction are nitrogen fixation, assimilatory nitrogen reduction, and denitrification. The major oxidation processes are nitrification and anaerobic ammonium oxidation (anammox). Each of these is described next in further detail. 

Figure 6.3 (Right) Schematic depiction of anammox cell showing the anammoxozome and nucleoid. (Left) Postulated pathway of anaerobic ammonium oxidation coupled to the ana-mmoxosome membrane resulting in a proton motive force and ATP synthesis via membrane-bound ATPases. HH, hydrazine hydrolase HZO, hydrazine oxidizing enzyme NIR, nitrite reductase. (Redrawn from van Niftrik etal., 2004 and Kuypers et al., 2006).

Recent work has demonstrated that denitrification is not the only pathway producing N2. The anaerobic ammonium oxidation (anammox) pathway has received considerable attention and appears to account for the lack of ammonium in some denitrifying regimes (e.g., Dalsgaard et al., 2003 Kuypers et al., 2003). In addition to to trace denitrification, parallel incubations with NH4+ are... 

Many other carboxylation reactions exist (Barton et al., 1991). For example, in methylo-trophic bacteria, formaldehyde and CO2 are combined to produce acetyl-CoA in the serine or hydroxypyruvate pathway. In contrast, the ribulose monophosphate cycle, which is another methylotrophic pathway of formaldehyde fixation, does not involve carboxylation steps. In addition to those described above, commonly found carboxylation reactions include those of pyruvate or phosphoenol pyruvate. In view of several relatively recent discoveries of novel CO2 assimilation pathways (e.g., the hydoxypro-pionate cycle and anaerobic ammonium oxidation) and growing interest in deep-subsurface microbiology, novel pathways of CO2 incorporation may be discovered in the near future. 

Nitrification-denitrification reactions are the major pathway for N loss from wetland ecosystems to the atmosphere. The process involves a series of sequential microbial processes that include mineralization of organic nitrogen to ammonium, oxidation of ammonium to nitrate, and denitrification of nitrate to nitrous oxide or dinitrogen gas. Nitrification-denitrification in wetlands occurs primarily in two zones the aerobic-anaerobic interface at the surface of the flooded soil or sediment, and the oxidized rhizosphere of wetland plants (Figure 16.6). 

The anaerobic and aerobic nitrifier denitrification pathways differ in that NO is an end product under anaerobic conditions rather than an intermediate compound. In addition, nitrogen dioxide-dependent NH3 oxidation by N. eutropha does not require ammonium monooxygenase (Schmidt et al., 2002), demonstrating that the two pathways are enzymatically different. In the absence of NH3, N. eutropha can use H2 or simple organic compounds as electron donors (Abeliovich and Vonhak, 1992 Bock et al., 1995). In contrast to the anammox process, which is strictly anaerobic, O2 does not inhibit N02-dependent NH3 oxidation and N2 production can occur even under aerobic conditions (Zart and Bock, 1998). However, Shrestha et al. (2002) observed N2 production... 

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