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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 />

<strong>International</strong> <strong>Seabed</strong> <strong>Authority</strong> 105

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