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Anammox - A Novel Microbial Process For Ammonium Removal: N. Shivaraman and Geetha Shivaraman
Anammox - A Novel Microbial Process For Ammonium Removal: N. Shivaraman and Geetha Shivaraman
Anammox1,2, a process of anaerobic Recently7 it was discovered that anam- moxasome is believed to be surrounded
ammonium oxidation, is an innovative mox makes a significant (up to 70%) by a bilayer membrane that consists of
technological advancement in the removal contribution to nitrogen cycling in the unique ‘ladderane’ lipids5.
of ammonia nitrogen in waste water. world’s oceans. Figure 1 shows the The structure and function of anam-
This new process combines ammonia and involvement of anammox process in the mox lipids has been published recently10.
nitrite directly into dinitrogen gas3. The biological nitrogen cycle including meta- Lipid membranes are essential to the
anammox reaction can be represented as bolic pathway for anaerobic ammonium functioning of cells facilitating the exis-
NH 4+ + NO 2− = N2 + 2H2O. This reaction oxidation. tence of concentration gradients of ions
is carried out by anammox bacteria be- Initially anammox research5 was fo- and metabolites. Microbial membrane
longing to the planctomycete group. Bac- cused on the basic properties of the proc- lipids are known to contain three-, five-,
teria capable of anaerobically oxidizing ess and on providing evidence for its six-, and even seven-membered aliphatic
ammonia were not identified earlier and microbial nature and the principles of the rings, but four-membered aliphatic cyclo-
were known as ‘lithotrophs missing from nitrogen and carbon metabolism. It butane rings have been reported for the
4
. Subsequently, this missing lith- appears that the anammox process is first time, in the dominant membrane
otroph was discovered1 and identified as based on energy conservation from an- lipids of two anammox bacteria. These
a new autotrophic member of the order oxic ammonium oxidation with nitrite as lipids contain up to five linearly fused
Planctomycetales. It appears that they the electron acceptor and hydrazine and cyclobutane moieties with cis ring junc-
might be the most primitive group of hydroxylamine as the intermediates. tions. These ‘ladderane’ moieties, rigid
bacteria, at the very root of the bacterial Carbon dioxide is used as the main car- and dense ladders of concatenated cyclo-
tree. Two of the anammox bacteria have bon source for growth. One of the key butane rings, occur in the membrane of
been named provisionally5: Candidatus enzymes of anaerobic ammonium oxida- the anammoxasome, the intracytoplasmic
‘Brocadia anammoxidans’ and Candida- tion is hydroxylamine oxidoreductase8. compartment, where anammox catabo-
tus ‘Kuenenia stuttgartiensis’. The former Its importance is illustrated by the fact lism takes place. A membrane with lad-
bacterium was responsible for anaerobic that it constitutes 10% of the total cell deranes in its core is highly impermeable
oxidation of ammonia observed in the protein. It catalyses the oxidation of hy- to passive diffusion of chemicals. Such a
Netherlands, while the latter organism drazine and hydroxylamine. The enzyme membrane is required to maintain con-
has been shown to be responsible for has been located in a membrane-boun- centration gradients during the very slow
ammonia oxidation anaerobically in sev- ded, ‘organelle’ named anammoxasome, anammox metabolism. It also protects
eral wastewater treatment plants in Ger- in the cytoplasm of the anammox cells9. the remainder of the cell from toxic
many and Switzerland. Interestingly the This ‘organelle’ appears to be the center anammox intermediates, viz. hydroxyl-
two species of the anammox bacteria of anaerobic ammonium oxidation. It is amine and hydrazine. Hydrazine and
which have been characterized are rarely quite likely that its function could be hydroxylamine were observed to diffuse
found together in a single anammox reac- containment of hydrazine. The anam- in and out of anammox cells as free
tor, they must have different, as-yet un- molecules3. The containment of these
known, ecological niches5. chemicals inside the anammoxasome was
It is well known that the biological considered impossible, since both com-
nitrogen cycle plays an important part in pounds diffuse through biomembranes.
the maintenance of nitrogen balance in The discovery10 of the unprecedented
global biosphere. The conventional bio- molecular structure of the anammox
logical nitrogen cycle involves microbial membrane lipids shows that the anam-
fixation of nitrogen gas to ammonia sym- moxasome membrane is much less per-
biotically and non-symbiotically which is meable than normal biomembranes
subsequently converted to organic nitro- because of the presence of unique ‘lad-
gen. The ammonia released from organi- derane’ lipids. From a bioenergetic per-
cally bound nitrogen and from man-made spective, the energy loss associated with
activities, is biologically oxidized aero- loss of one molecule of hydrazine from
bically to nitrite and then to nitrate. The the anammox cell is equivalent to at least
resultant nitrate and nitrite are reduced to 10 full catabolic cycles, leading to 50%
nitrogen gas by a denitrifying group of decrease in biomass yield at 10% hydra-
bacteria using some electron donors (or- zine loss. For these reasons, limitation of
ganic or inorganic compounds). It has diffusion is very important for these bac-
been indicated6 that anaerobic ammonia Figure 1. Involvement of anammox pro- teria which are slow growers. One of
cess in biological nitrogen cycle including
oxidation also could be a major biologi- metabolic pathway for anaerobic ammo- these lipids was identified as a sn-2-gly-
cal activity to be included in the nitrogen nium oxidation (adapted from refs 3 and cerol monoether with a C20 alkyl chain
cycle, thus necessitating its modification. 5). containing four rings. The second lipid