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The ecology of eelgrass meadows in the Pacific Northwest: A ...

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total annual primary prduction and 14% <strong>of</strong><br />

<strong>the</strong>ir total excreted material <strong>in</strong>to <strong>the</strong><br />

estuar<strong>in</strong>e system.<br />

3.4 ORGANIC AND INORGANIC NUTEUENI'<br />

CYCLING<br />

<strong>The</strong> anatomy <strong>of</strong> <strong>eelgrass</strong>, like that <strong>of</strong> all<br />

o<strong>the</strong>r seagrasses, is modified for<br />

metabolism, growth, and reproduction <strong>in</strong><br />

<strong>the</strong> sea.<br />

1. <strong>The</strong> cut<strong>in</strong> layer over <strong>the</strong> leaf is<br />

th<strong>in</strong>.<br />

2. Leaf blades are flattened and th<strong>in</strong><br />

with a very high surf ace-to-volume<br />

ratio.<br />

3. <strong>The</strong>re is an extensive lacuna1 system<br />

which gives buoyancy (<strong>the</strong>y store and<br />

transport a great quantity <strong>of</strong> O2 and<br />

a2'<br />

4. Chloroplasts are densely packed <strong>in</strong><br />

<strong>the</strong> epidermal layer suround<strong>in</strong>g<strong>the</strong><br />

leaf.<br />

5. <strong>The</strong>re is an aerenchyma composed <strong>of</strong><br />

large, th<strong>in</strong>-walled cells which adjo<strong>in</strong><br />

<strong>the</strong> lacunae and facilitate gas and<br />

solute diffusion.<br />

6. <strong>The</strong>re is a reduced amount <strong>of</strong><br />

mechanical support, allow<strong>in</strong>g <strong>the</strong><br />

leaves to f Lex and bend on an ebb<strong>in</strong>g<br />

tide, which ma<strong>in</strong>ta<strong>in</strong>s <strong>the</strong>ir wetness<br />

and guarantees expos<strong>in</strong>g as much<br />

photosyn<strong>the</strong>tic surface to solar<br />

radiation as possible. At <strong>the</strong> same<br />

time <strong>the</strong> leaves are strong enough to<br />

resist break<strong>in</strong>g when <strong>the</strong>y are whipped<br />

dur<strong>in</strong>g wave action e erg us on et al.<br />

1980; Zieman 1982) .<br />

<strong>The</strong> major problem with procurement <strong>of</strong><br />

certa<strong>in</strong> nutrients, such as carbon, by<br />

seagrasses is that rates <strong>of</strong> gaseous<br />

diffusion <strong>in</strong> water are several orders <strong>of</strong><br />

magnitude lower than <strong>in</strong> air. Also, when<br />

<strong>the</strong> pH <strong>of</strong> seawater rises dur<strong>in</strong>g very<br />

active photosyn<strong>the</strong>sis (from 8.2 to 8.9 or<br />

higher), free C02 <strong>in</strong> <strong>the</strong> water is greatly<br />

reduced or becomes unavailable, as does<br />

<strong>the</strong> KO3-ion. It is now known that<br />

seagrasses obta<strong>in</strong> most <strong>of</strong> <strong>the</strong>ir nutrients<br />

from <strong>the</strong> sediments, which ma<strong>in</strong>ta<strong>in</strong> a much<br />

lower pH <strong>in</strong> <strong>the</strong> deep anoxic layer beneath<br />

<strong>the</strong> very th<strong>in</strong> surface oxic zone.<br />

Numerous papers have documented <strong>the</strong> uptake<br />

<strong>of</strong> nutrients (carbon, phosphorus, and<br />

nitrogen, <strong>in</strong> particular) from <strong>the</strong><br />

sediments by <strong>the</strong> root system,<br />

translocation through <strong>the</strong> seagrass plant,<br />

and release to <strong>the</strong> epiphytes and water<br />

column. <strong>The</strong> plants and epiphytes can also<br />

absorb <strong>the</strong>se nutrients from <strong>the</strong> water to<br />

release <strong>the</strong>m to <strong>the</strong> <strong>in</strong>terstitial water <strong>in</strong><br />

<strong>the</strong> sediments (MCRO~ and Barsdate 1970;<br />

Mcmy et al. 1972; McWy and Wr<strong>in</strong>g 1974;<br />

Harl<strong>in</strong> 1975, 1980; Penhale and 3nith 1977;<br />

Wtzel and Penhale 1979; Penhale and<br />

Thayer 1980; Short 1981) . All <strong>the</strong> available<br />

work done states that sediments are<br />

<strong>the</strong> preferred source for nutrient uptake.<br />

<strong>The</strong>se laboratory experiments were<br />

re<strong>in</strong>forced by recent field experiments<br />

us<strong>in</strong>g fertilizers. Orth (197733) used<br />

Osmacote (14-14-14; a slow-release<br />

fertilizer), placed <strong>in</strong> <strong>the</strong> sediments, and<br />

observed dramatic <strong>in</strong>creases <strong>in</strong> plant<br />

stand<strong>in</strong>g crops. Eelgrass root/rhizomes<br />

<strong>in</strong>creased 30%, <strong>the</strong> leaf crop was three to<br />

four times higher, and shoot density<br />

<strong>in</strong>creased. Increases <strong>in</strong> <strong>the</strong> stand<strong>in</strong>g crop<br />

were also observed follow<strong>in</strong>g fertilization<br />

<strong>of</strong> <strong>eelgrass</strong> <strong>in</strong> mode Island (Harl<strong>in</strong> and<br />

Thorne-Miller 1981) .<br />

Even though <strong>eelgrass</strong> needs a variety <strong>of</strong><br />

macro- and micronutrients, most <strong>of</strong> <strong>the</strong><br />

research has concentrated on carbon,<br />

phosphorus, and nitrogen. Phosphorus<br />

supply does not appear to be limit<strong>in</strong>g. In<br />

Alaska, McRoy and Barsdate (1970) found<br />

that <strong>eelgrass</strong> plants were a phosphorus<br />

pump from <strong>the</strong> sediments to <strong>the</strong> water<br />

column, but <strong>in</strong> North Carol<strong>in</strong>a <strong>the</strong> roots<br />

reta<strong>in</strong>ed most <strong>of</strong> <strong>the</strong>ir absorbed phosphorus<br />

(Penhale and Thayer 1980). McRoy et al.<br />

(1972) estimated that phosphorus turnover<br />

times <strong>in</strong> <strong>the</strong> <strong>eelgrass</strong> <strong>meadows</strong> <strong>in</strong> Izembek<br />

Lagoon, Alaska, vary from two turnovers<br />

per year to one every 2 years.<br />

Eelgrass has three sources <strong>of</strong> <strong>in</strong>organic<br />

carbon (C02, HCO~-) : recycled from<br />

respiration and p otorespiration, <strong>the</strong><br />

water column (which is not a likely source<br />

dur<strong>in</strong>g active photcsyn<strong>the</strong>sis, ow<strong>in</strong>g to an

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