72 4 Taxonomy, Physiology, and Ecology of Aquatic Microorganisms
The anammox bacteria are characterized by several
striking properties:
(a) They all possess one anammoxosome, a membrane
bound compartment inside the cytoplasm which is
the site of anammox catabolism.
(b) Further, the membranes of these bacteria mainly
consist of ladderane lipids which are rare in living
organisms.
(c) Hydrazine (normally used as a highenergy rocket
fuel, and poisonous to most living organisms) is a
byproduct of these organisms.
(d) Finally, the organisms grow very slowly, the gen
eration or doubling time being nearly 2 weeks.
The anammox process was originally found to
occur only from 20°C to 43°C, but more recently, ana
mmox has been observed at temperatures from 36°C to
52°C in hot springs and 60°C to 85°C at hydrothermal
vents located in the ocean floor.
Reduction under anaerobic conditions can also occur
through anaerobic ammonia oxidation (Anammox) thus:
Because denitrifying bacteria are principally het
erotrophic, in some wastewater treatment plants, small
amounts of methanol are added to the wastewater to
provide a carbon source for the bacteria.
Nitrogen fixation, nitrification, and denitrification
are interlinked in the nitrogen cycle. The nitrogen
cycle in the marine environment is given in Fig. 4.1 5.
The Sulfur Cycle in the Aquatic System and
Bacteria
In the environment, through changes brought about
mostly by bacteria, sulfur changes from one form to
the other: From hydrogen sulfide (H 2 S) to sulfate via
elemental sulfur (S^0 ) and sulfate is changed again to
hydrogen sulfide. Hydrogen sulfide is also evolved
from hot springs and volcanoes, and occurs when dead
animals, the excreta of animals, and dead plants are
decomposed by bacteria. The compound is oxidized to
sulfuric acid by the sulfuroxidizing bacteria and pho
tosynthetic sulfur bacteria via elemental sulfur. The
change of hydrogen sulfide to elemental sulfur occurs
also abiotically in the presence of molecular oxygen.
Dimethyl sulfide (CH 3 ) 2 S is produced by marine
algae and marine cyanobacteria and contributes to the
typical smell of thee sea. Dimethyl sulfide is degraded
by bacteria such as Thiobacillus and Hyphomicrobium,
leading to the formation of acid. The various transfor
mations are summarized in Fig. 4.1 6. A major group of
bacteria important in the global economy of sulfur,
especially in aquatic environments are the sulfate reduc
ing bacteria. They will be discussed briefly below.
The sulfate reducing bacteria (SRB) are ubiquitous
anaerobes found in diverse environments. They include
several groups of bacteria that use sulfate as an oxidiz
ing agent, reducing it to sulfide (Fig 4.1 7 ) (Luptakova
2007 ). They can also utilize other sulfur compounds,
including sulfite, thiosulfate, and elemental sulfur in
a type of metabolism known as dissimilatory, because
NH 4 NO2 22N 2H O.
+−+ →+
Oxidation
state
Anammox
Assimilation
Nitr
ification
Assimilation
(NH 3 )NORG N H 3
N 2 O
N 2
(X) (Y)
+V
+IV
+III
−III
+II
−II
+I
−I
0
N 2 O
NO 2 − NO 2 − NO 2 −
NO 3 −
Catabolism
Denitr
ification
Nitrogen Fixation
Fig. 4.15 Nitrogen cycle
in the marine environment
(From Codispoti et al. 2001.
With permission)
“X” and “Y” are intracel
lular intermediates that do
not accumulate in water
column