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3.1 EPR Vent Ecosystems<br />
A schematic representation of the general appearance of vent ecosystems on the<br />
northern EPR is shown in Figure 3. More vent species are known from sites on the<br />
northern EPR than from any other spreading ridge on the planet. This may in part be due<br />
to the longer history of research in the area, but there are legitimate theoretical<br />
arguments that relate this greater biodiversity to the history of seafloor spreading and the<br />
abundance of vent habitat along the northern EPR 9,10. Chemosynthetic microbial growth<br />
provides the primary production of biomass, and occurs in three distinctly different<br />
habitats: endosymbioses, ectosymbioses, and free-living on animal and mineral surfaces.<br />
Endosymbioses<br />
There are three general models for the functioning of the most common forms of<br />
host-symbiont associations at EPR vents: vestimentiferan worms, clams and mussels.<br />
Microbial symbionts directly nourish their hosts through lysis (digestion of symbionts<br />
within host cells) or through secretion of organic matter that is subsequently absorbed by<br />
host tissues 4.<br />
Type 1 - The most evolved symbiosis occurs in the vestimentiferan tube worms -<br />
which have no mouth or digestive system, and are entirely reliant on their symbiotic<br />
bacteria for nutrition. In the tubeworms the symbiotic bacteria are housed in a<br />
specialized organ known as the trophosome. Substrates for microbial metabolism (HS - ,<br />
CO2, O2, etc) are taken up at the gills and transported to the trophosome by the worm's<br />
blood 4. Physiological and biochemical aspects of this symbiosis have been extensively<br />
studied in the vestimentiferan Riftia pachytila 4.<br />
Type 2 - Vesicomyid clams found at vent sites are filter-feeding animals, but<br />
their digestive tract is highly reduced and experiments have shown that they are unable<br />
to survive without a supply of hydrogen sulphide for their symbionts. They host their<br />
symbionts in the tissue of their large modified gills. CO2 and O2 diffuse directly into the<br />
gills from the external environment, while the clams take up H2S into their blood through<br />
their foot, which they extend into fractures, or sediments where there is diffuse<br />
hydrothermal flow 4. The large size attained by the giant vent clam Calyptogena magnifica<br />
attests to the productive nature of this symbiosis.<br />
Type 3 - Like the clams, the EPR vent mussel Bathymodiolus thermophylus also<br />
houses its symbionts in gill tissue, is a filter feeder and has a functional digestive system.<br />
Unlike the clams, filtration of particles of organic matter from the surrounding water<br />
appears to provide a significant supplement to the mussel's nutrition. When<br />
experimentally moved away from active venting these mussels do survive, although<br />
considerable weight loss occurs. The mussel blood lacks specific proteins for the binding<br />
and transport of sulphide and oxygen that are found in the vestimentifera and clams 4.<br />
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