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

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involved processes). It is thus appropriately described by a 'transitional'<br />

<strong>model</strong>. However, by fitting data, only <strong>the</strong> shape that is matched or not by<br />

<strong>the</strong> <strong>model</strong> is considered. Consequently, 'dimensional' <strong>model</strong>s fit equally<br />

well, although <strong>the</strong>ir ma<strong>the</strong>matical formulation does not match biological<br />

observation.<br />

Considering this distinction, we can now formulate a <strong>model</strong> that is both<br />

'dimensional' and 'transitional':<br />

⎛ −k1⋅t' 1−e m<br />

⎞<br />

0 ∞ −k2⋅t' Yt (') = Y+∆Y⎜ ⎟<br />

⎝1+ l ⋅e<br />

⎠<br />

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

(40)<br />

Y being any kind <strong>of</strong> measurement <strong>of</strong> size and m (corresponding to d in <strong>the</strong><br />

Richards <strong>model</strong>) indicating <strong>the</strong> power transformation required to be in <strong>the</strong><br />

best 'dimension <strong>of</strong> growth'. To fur<strong>the</strong>r generalize eq. 40, we could also<br />

replace k1·t' (<strong>the</strong> chronological time modulated by a constant kinetic<br />

parameter) with tM, <strong>the</strong> metabolic –or physiologic– time (Brody, 1937),<br />

using:<br />

t = f( t, x , x ,..., x )<br />

(41)<br />

M 1 2 n<br />

where x1-n are environmental variables that modulate growth, e.g., <strong>sea</strong>son<br />

(Cloern & Nichols, 1978) or temperature (Muller-Feuga, 1990). This<br />

gives:<br />

−t<br />

m<br />

M ⎛ 1−e ⎞<br />

= 0 +∆ ∞ ⎜ −kt ⋅ ⎟ M<br />

Yt (') Y Y<br />

⎝1+ l ⋅e<br />

⎠<br />

(42)<br />

where k = k2/k1. This is a general functional <strong>model</strong> for an asymptotic<br />

growth that derives from <strong>the</strong> von Bertalanffy 1 curve. Therefore, we call it<br />

a generalized von Bertalanffy <strong>model</strong>. This <strong>model</strong> is impossible to fit<br />

without some precautions because <strong>the</strong> two effects, 'dimension' (m) and<br />

'transition' (l and k), are impossible to separate with solely a shape<br />

criterion. From a fitting point <strong>of</strong> view, this <strong>model</strong> is overparameterized<br />

(Draper & Smith, 1998). The dimension parameter m must first be<br />

175

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