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Principles of terrestrial ecosystem ecology.pdf

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230 10. Aquatic Carbon and Nutrient Cycling<br />

Depth (m)<br />

20<br />

60<br />

100<br />

140<br />

0.1 1 10 100 0.1 1 10 100<br />

0.1 1 10 100<br />

0<br />

0<br />

G<br />

B<br />

Y<br />

R<br />

tively low light intensity, from 5 to 25% <strong>of</strong><br />

full sun, depending on the algal group (Valiela<br />

1995). Photosynthesis declines at higher light<br />

levels, so maximum photosynthesis usually<br />

occurs at about 10m depth on sunny days. One<br />

reason that phytoplankton function as shade<br />

plants is that they mix vertically through the<br />

water column, so an individual cell spends relatively<br />

little time near the surface. Photosynthetic<br />

acclimation to high light requires about<br />

12h, which is probably longer than the time that<br />

most phytoplankton would be exposed to high<br />

light. In the ocean and clear lakes, UV-B radiation<br />

may also contribute to low photosynthetic<br />

rates in surface waters, raising questions about<br />

the consequences <strong>of</strong> ozone holes and increased<br />

UV-B at high latitudes. Light appears to limit<br />

ocean and lake production at the water surface,<br />

primarily beneath ice cover or during winter at<br />

high latitudes when the sun angle is low. At<br />

depth, light limits production in all aquatic<br />

habitats.<br />

Primary producers exhibit a greater diversity<br />

<strong>of</strong> pigment types in the ocean than on land,<br />

presumably related to the complex light<br />

environment <strong>of</strong> water. Red algae, which are<br />

abundant at depth in tropical oceans, and<br />

5<br />

10<br />

15<br />

R<br />

20<br />

25<br />

30<br />

B G<br />

W<br />

10<br />

20<br />

B G<br />

Oceanic water Coastal water Forest<br />

Figure 10.5. Light quality at different depths in<br />

ocean and coastal waters and in forests (Chazdon<br />

and Fetcher 1984a). R, red; Y, yellow; G, green; B,<br />

Irradiance (% <strong>of</strong> incident light) [log scale]<br />

blue; W, white. (Modified with permission from<br />

Springer-Verlag; Valiela 1995.)<br />

brown algae like kelp, which are abundant at<br />

depth in cool temperate oceans, have pigments<br />

that absorb the blue light that is<br />

available to them. Surface algae tend to have<br />

green pigments. These differences in algal pigments<br />

may also be adaptations to light quantity<br />

per se.<br />

Reduced sulfur compounds provide the<br />

energy for carbohydrate synthesis from CO2<br />

in some anaerobic aquatic habitats. Although<br />

most organisms depend on carbon fixed<br />

through light-dependent photosynthesis, some<br />

bacteria use the energy <strong>of</strong> reduced sulfur compounds<br />

to reduce CO2 to form organic compounds.<br />

Entire <strong>ecosystem</strong>s are built on such a<br />

base near hydrothermal vents in zones <strong>of</strong> seafloor<br />

spreading in midocean regions. Although<br />

hydrothermal vents account for only a tiny fraction<br />

<strong>of</strong> total ocean production, they support<br />

unique communities and complex food webs<br />

that are completely independent <strong>of</strong> energy<br />

input from the sun (Karl et al. 1980). Similar<br />

chemosynthesis occurs in anaerobic sediments,<br />

but these sulfur-dependent oxidation–reduction<br />

reactions usually account for only a small<br />

fraction <strong>of</strong> the total carbon budget <strong>of</strong> these<br />

environments.<br />

R

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