Bacteria living in the
first few
millimeters to meter of sediment live via chemoautrophic reactions
after all
available oxygen has been utilized (eg. Jorgensen 1982, Ravenschlag et al. 2000).
First, nitrate is reduced, then manganese and next iron are
oxidized. However, less is known about
the energy sources of bacteria the deeper one moves in the sediment
column. Types of bacteria identified thus
far,
categorized by their chemoautotrophic energy source include: aerobic
ammonifers; nitrate-reducers, both to ammonia and nitrogen;
fermentative
anaerobic heterotrophs; sulfate reducers, including acetate, lactate,
and
methane utilizers that produce CO2 from CH4 in
initial
enrichments but for which activity was not sustained on subcultures;
methanogens; acetogens; and anaerobic hexadecane oxidizers (Parkes et
al. 2000).
As mentioned in the
introduction, sulfate reduction was
long considered to be the final step in the reduction of organic
material as
one moved down the sediment column.
However, it appears that the “last step” of
chemoautotrophic or
chemosynthetic life in the sediments may change depending on a variety
of
factors and therefore be different from locality to locality. D’Hondt et al. 2002 claimed that SO42
reduction, methanogenesis, and
fermentation are the principal degradative metabolic processes
in
subsurface (>1.5 mbsf) marine sediments for three reasons:
First,
at the sediment-water interface, concentrations of dissolved
SO42
are more than 50 times as
great as concentrations of all electron acceptors with
higher
standard free energies combined. Second, external electron acceptors,
such as
iron and manganese, which yield more energy than SO42
typically disappear within the
first few centimeters to tens of meters of sediment depth. Thirdly, once all SO42
has been reduced, methanogenesis
and fermentation are the principal remaining avenues of
metabolic
activity (D'Hondt et al. 2002).
Methanogenesis
can be achieved
through two different reactions: H2/CO2
where HCO3– is reduced to CH4,
and acetate
oxidation (Wellsbury and Parkes 2000, Parkes et al. 2000).
It is believed t sulfate
reducing bacteria can out compete methanogens for organic carbon down
to
sulfate concentrations of about 3mM.
Below this concentration, methanogenesis dominates.
However, methane can be used in sulfate
reduction as well. These organisms keep
methane concentrations low through the sediment column in comparison to
sulfate. At two gas hydrate zones, the
Peru Margin and the