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Growth model of the reared sea urchin Paracentrotus ... - SciViews

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considered factor on <strong>the</strong> curve's shape). For instance, current <strong>model</strong>s<br />

hardly cope with a superimposed effect <strong>of</strong> intraspecific competition on<br />

growth, though <strong>the</strong> latter was evidenced in <strong>sea</strong> <strong>urchin</strong>s (Ebert, 1977;<br />

Himmelman, 1986; Levitan, 1988; Grosjean et al, 1996) as in many o<strong>the</strong>r<br />

species (Branch, 1974, for <strong>the</strong> limpet Patella cochlear Born; Timmons &<br />

Shelton, 1980, for <strong>the</strong> largemouth bass Micropterus salmoides (Lacepede);<br />

Kautsky, 1982, for <strong>the</strong> mussel Mytilus edulis L.).<br />

One method to <strong>model</strong> growth in a functional way is by building<br />

equations that directly represent underlying processes, i.e., by elaborating a<br />

dynamic <strong>model</strong> that balances inputs (food, oxygen intake…) and outputs<br />

(carbon dioxide, feces…) from which it is possible to calculate variation <strong>of</strong><br />

size with time, thus yielding an estimation <strong>of</strong> growth. This is <strong>the</strong><br />

bioenergetic and/or ecophysiologic approach, using <strong>the</strong> scope for growth<br />

concept, which has proved very successful for filter feeders (Willows,<br />

1992). Such an approach requires a lot <strong>of</strong> measurements and equations. It<br />

is most <strong>of</strong>ten used in <strong>the</strong> simplest cases, where environmental conditions<br />

are constant, or vary in a very predictable way, like in a protected reef<br />

lagoon for cultivated pearl oysters (Pouvreau et al, 2000). Indeed, similar<br />

studies on European oyster cultures in <strong>the</strong> intertidal zone –a very changing<br />

and unpredictable environment– lead to a much more complex <strong>model</strong><br />

(Bacher et al, 1991). As far as we know, no such <strong>model</strong> has been<br />

completely successful when applied to <strong>sea</strong> <strong>urchin</strong>s because a large part <strong>of</strong><br />

<strong>the</strong> carbon or energy absorbed is lost as dissolved organic matter that is<br />

hard to quantify and to enter in equations (Miller & Mann, 1973; Lawrence<br />

& Lane, 1982).<br />

Being a basic feature <strong>of</strong> life, growth has been widely explored but it<br />

still remains unsatisfactorily <strong>model</strong>led in a functional way, possibly<br />

because <strong>of</strong> <strong>the</strong> approach used. Most growth <strong>model</strong>s were elaborated from<br />

<strong>the</strong>ir differential equations. Functions obtained by solving <strong>the</strong>se equations<br />

were <strong>the</strong>n systematically used to determine growth <strong>of</strong> a "mean individual"<br />

(by least-square regression) and no parameter <strong>of</strong> <strong>the</strong> <strong>model</strong> was<br />

constrained using particular knowledge (such as size at birth or at<br />

Part IV: A growth <strong>model</strong> with intraspecific competition<br />

140

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