Subsequently,
in order to evaluate in more detail the community
composition, the authors used fulllength 16S ribosomal RNA gene clone
libraries constructed from basaltic lavas and surrounding sea water
samples. These phylogenetic analyses revealed that both the
basalt-hosted biospheres (EPR and Hawaii), were harbouring
high-richness bacterial communities and that community membership was
shared between these sites. A statistical approach was then used to
evaluate the species richness (number of operational taxonomic units)
as compared to other oceanic environments analyzed in other studies
(e.g. an hydrothermal white smoker, the upper water column of the
Sargasso Sea, hydrothermal fluids from the Mid-Atlantic Ridge and
deep-subsurface sediments from the Nankai Trough). These comparative
analysis, revealed that abundance, phylogenetic diversity and
richness of Bacteria in these other deep-sea environments were
clearly lower and much different than EPR and Hawaii deep-sea basalts. The 21 taxonomic groups recovered from basalt were dominated
by Proteobacteria (68% and
66% of all sequences in EPR and Hawaii respectively), while
non-Proteobacteria groups included Plantomycetes (8%/5%),
Actinobacteria (7%/8%), Bacteroidetes (4%/1%),
Acidobacteria (3%/4%) and Verrucomicrobia (2%/2%). Interestingly, the OTU richness for the two geographically separated
basalt communities (EPR and Hawaii) showed considerable overlap in
community membership, suggesting that oceanic basalt microbes are
widely distributed among this biotope.
These
differences in phylogenetic diversity, species richness, and total
biomass between the basaltic lavas and overlying sea water raised
questions about what energy source fuel this biosphere. Potential
energy sources capable of sustaining microbial life in ocean crust
include hydrothermal input of manganese and
iron (chemolithoautotrophic growth) and dissolved organic carbon in
sea water or hydrothermal fluids (heterotrophic growth). However,
according to the authors, the most plausible explanation is that
oceanic lithosphere exposed at the sea floor undergoes seawater-rock
alteration reactions and these reactions are capable of supplying
sufficient energy for chemolithoautotrophic microbial growth. Lava
surfaces in fact, are composed predominantly of volcanic glass, a
highly reactive rock component that contains reduced elemental
species such as iron, sulphur and manganese. Oxygen and nitrate in
deep sea water oxidize these constituents and
chemolithoautotrophic microorganisms can potentially exploit the free
energy changes associated with these redox reactions for their
metabolic requirements. Laboratory studies have already demonstrated
that iron-oxidizing bacteria isolated from the sea floor are able to
use rock and minerals, including glassy basalt, for metabolism and
growth. The authors estimated that about 6x107-6x109
cells
per g basalt may be supported through these reactions and actually,
cell densities in EPR basalts were falling
exactly within this range. So in conclusion,
alteration reactions in the upper ocean crust may fuel microbial
ecosystems at the sea floor, which constitute a trophic base of the
basalt biotope, with important implications for deep-sea carbon
cycling and chemical exchange between basalt and sea water. This
hypothesis supports the understanding of the phylogenetically rich
and distinct nature of the basalt biotope. The enrichment of taxa
from diverse metabolic groups may result from the establishment of
chemical microenvironments within or on rock cavities and surfaces
during alteration, mineral precipitation and biofilm formation. This
niche creation would allow for a greater variety of redox reactions
and metabolic pathways (e.g. heterotrophic, anaerobic, or reductive)
including those supporting complex organotrophic and mixotrophic
communities.
Reference:
Santelli, C.M., B.N. Orcutt, E.
Banning, W. Bach, C.L. Moyer, M.L. Sogin, H. Staudigel, and K.J.
Edwards. (2008). Abundance and diversity of microbial life in ocean
crust. Nature 453:653-656.
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