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

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CHAPTER 2<br />

THE BIOLOGY OF EELGRASS<br />

2.1 VEGETATIVE GlXWlTd STRATEGIES<br />

Sauvageau (1889, 1890, 1891), Setchell<br />

(19291, Phillips (1972), and Toml<strong>in</strong>son<br />

(1974, 1980) described <strong>the</strong> morphology and<br />

anatomy <strong>of</strong> <strong>eelgrass</strong>. Eelgrass leaves<br />

possess an anatomy typical <strong>of</strong> submerged<br />

hydrophytes: namely, cut<strong>in</strong> on <strong>the</strong> leaf is<br />

th<strong>in</strong>, <strong>the</strong>re are no stomata, chloroplasts<br />

are densely packed and lie pr<strong>in</strong>cipally <strong>in</strong><br />

<strong>the</strong> epidermis, <strong>the</strong> <strong>in</strong>ternal cellular<br />

structure consists <strong>of</strong> large th<strong>in</strong>-walled<br />

aerenchyma cells with numerous air canals<br />

(lacunae), and <strong>the</strong> vascular tissue has<br />

reduced xylem (Toml<strong>in</strong>son 1980). As with<br />

grass-like monocots whose leaves stand<br />

erect, <strong>the</strong>re are no dorsal or ventral<br />

sides to <strong>the</strong> leaf blade. <strong>The</strong> lacunae are<br />

cont<strong>in</strong>uous from <strong>the</strong> leaf blade through <strong>the</strong><br />

rhizome to <strong>the</strong> root tips and carry o2 and<br />

C02 throughout <strong>the</strong> plant. Because <strong>the</strong>se<br />

lacunae may also store and recycle 02, <strong>the</strong><br />

standard method <strong>of</strong> measur<strong>in</strong>g prlmary<br />

productivity by measur<strong>in</strong>g O2 changes <strong>in</strong><br />

<strong>the</strong> water around <strong>eelgrass</strong> plants is not<br />

accurate (Hartmanand Brown1967; Jacobs<br />

1979; Zieman and Wetzel 1980). <strong>The</strong> leaftips<br />

are rounded, while <strong>the</strong> blades are<br />

straplike. Leaf width depends on <strong>the</strong><br />

severity <strong>of</strong> <strong>the</strong> climate <strong>in</strong> a region, <strong>the</strong><br />

season, and tidal zone (Setchell 1920,<br />

1929; Phillips 1972; Kentula 1983). <strong>The</strong><br />

leaves are produced by a meristem term<strong>in</strong>al<br />

on a short shoot (erect branch from <strong>the</strong><br />

horizontal rhizome). <strong>The</strong> oldest shoot<br />

(def<strong>in</strong>ed as an erect branch with a bundle<br />

<strong>of</strong> leaves) is term<strong>in</strong>al on <strong>the</strong> rhizome.<br />

Occasionally <strong>the</strong> meristem on <strong>the</strong> term<strong>in</strong>al<br />

shoot gives rise to a lateral rhizome<br />

branch with a meristem that produces<br />

leaves.<br />

Depend<strong>in</strong>g on <strong>the</strong> length <strong>of</strong> <strong>the</strong> grow<strong>in</strong>g<br />

season, <strong>the</strong> number <strong>of</strong> leaves produced <strong>in</strong> a<br />

year, and<strong>the</strong> number <strong>of</strong> lateral branches<br />

produced, <strong>the</strong> <strong>in</strong>dividual <strong>eelgrass</strong> plant<br />

has a variable growth and expansion rate<br />

over <strong>the</strong> bottom. <strong>The</strong>re is one rhizome<br />

ncde prcduced for each new leaf <strong>in</strong>itiated;<br />

<strong>the</strong> same meristem which produces a leaf<br />

also produces a new <strong>in</strong>ternode for <strong>the</strong><br />

elongat<strong>in</strong>g branch. Setchell (1929)<br />

<strong>the</strong>orized that <strong>eelgrass</strong> plants produced<br />

two lateral branches dur<strong>in</strong>g a grow<strong>in</strong>g<br />

season and that <strong>the</strong> term<strong>in</strong>al shoot always<br />

flowered dur<strong>in</strong>g <strong>the</strong> second year follow<strong>in</strong>g<br />

development from a seed. In this scheme<br />

<strong>eelgrass</strong> is a biennial plant.<br />

Eelgrass rhizome are buried from 3-4 cm<br />

(1.2-1.6 <strong>in</strong>ches) up to 20 cm (8.0 <strong>in</strong>ches)<br />

deep <strong>in</strong> sediment, depend<strong>in</strong>g on <strong>the</strong><br />

sediment consistency. In firmer<br />

substrates, rhizomes may be only half as<br />

deep as <strong>in</strong> s<strong>of</strong>t muddy substrates. Two<br />

bundles <strong>of</strong> roots are produced at each<br />

rhizome node.<br />

New leaves grow on alternat<strong>in</strong>g sides from<br />

<strong>the</strong> meristem on <strong>the</strong> shoot. Subtidal<br />

shoots <strong>in</strong> Puget Sound typically carry five<br />

leaves each dur<strong>in</strong>g summer and four <strong>in</strong><br />

w<strong>in</strong>ter e hilli ips 1972), while <strong>in</strong>tertidal<br />

shoots <strong>in</strong> Oreyon averaged four leaves each<br />

<strong>in</strong> June and as few as 2.5 each <strong>in</strong> August<br />

(Kentula 1983). Keller and Harris (1966)<br />

reported that <strong>the</strong> number <strong>of</strong> leaves per<br />

shoot <strong>in</strong> Humboldt Bay ranged from two to<br />

thirteen but averaged three to four.<br />

Kentula (1983) analyzed <strong>the</strong> growth <strong>of</strong> a<br />

leaf <strong>in</strong> relation to its age-position on<br />

<strong>the</strong> shoot. <strong>The</strong> youngest leaf was<br />

designated No. 1. <strong>The</strong> greatest proprtion<br />

<strong>of</strong> growth occurred <strong>in</strong> position No. 2. In<br />

April-May <strong>the</strong>se leaves accounted for 48%-<br />

65% <strong>of</strong> <strong>the</strong> total growth <strong>of</strong> <strong>the</strong> shoot and<br />

from 75%-95% <strong>of</strong> <strong>the</strong> growth from June-<br />

October.

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