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

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Set L describes <strong>the</strong> largest size reached by <strong>sea</strong> <strong>urchin</strong>s at maximum<br />

growth speed with time. P. lividus having a determinate or asymptotic<br />

growth (see Fig. 28A), final increase <strong>of</strong> size ∆D∞ = D∞ – D0 is finite for<br />

t' → ∞. However, maximum size cannot be reached instantaneously. If <strong>the</strong><br />

largest individuals in <strong>the</strong> actual dataset are not inhibited at all, <strong>the</strong>y can be<br />

used as a reference for <strong>the</strong> whole cohort to define this maximum growth<br />

curve. Supposing that a von Bertalanffy 1 curve best fit <strong>the</strong> largest fraction<br />

<strong>of</strong> <strong>the</strong> size distributions (see next section), it is an appropriate <strong>model</strong> for set<br />

L. Because we use <strong>the</strong> t' time-scale here, we choose a parameterization <strong>of</strong><br />

<strong>the</strong> <strong>model</strong> such as D'(t' = 0) = D0:<br />

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

−k1⋅t' Dt' ( ) = D +∆D⋅(1− e )<br />

(25)<br />

0<br />

∞<br />

Membership to <strong>the</strong> L set with time, ML(t'), is <strong>model</strong>led with a logistic<br />

function (a classical <strong>model</strong> for a transition in fuzzy sets, Cox, 1999)<br />

(Fig. 30):<br />

1<br />

M ( t')<br />

=<br />

1+ l ⋅e<br />

L −k2⋅t' (26)<br />

Membership to <strong>the</strong> S set with time, MS(t'), is complementary so that MS<br />

and ML add up to one:<br />

1<br />

M ( t') = 1 − M ( t')<br />

= 1− 1+ l ⋅e<br />

S L −k2⋅t' (27)<br />

Thus, full-growing individuals belong to <strong>the</strong> L set from <strong>the</strong> beginning. The<br />

stronger <strong>the</strong> growth inhibition, <strong>the</strong> longer o<strong>the</strong>r individuals remain in <strong>the</strong> S<br />

set before gradually shifting to <strong>the</strong> L set. The fuzzy <strong>model</strong> integrates <strong>the</strong><br />

effect <strong>of</strong> intraspecific competition (or any o<strong>the</strong>r inhibition mechanism<br />

having a similar effect on growth) as a delayed transition from S to L sets,<br />

as explicitly quantified by parameter l (<strong>the</strong> lag or position <strong>of</strong> <strong>the</strong> inflexion<br />

point in <strong>the</strong> membership curves, see Fig. 30). If l = 0, <strong>the</strong>re is no inflexion<br />

point and ML(t') = 1; growth occurs at maximum speed from start and<br />

follows set L, that is, a von Bertalanffy curve. While parameter k1<br />

151

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