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

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tend to form aggregates where a pH gradient is measurable from outside to<br />

inside. Smaller animals are more likely found inside and larger <strong>sea</strong> <strong>urchin</strong>s<br />

outside. This was described as a protective behavior against predators in<br />

<strong>the</strong> field (Tegner & Dayton, 1977; Tegner & Levin, 1983; Levitan &<br />

Genovese, 1989; Ebert, 1998). But <strong>the</strong>n, parallel gradients in both pH and<br />

size distribution could be <strong>the</strong> explanation <strong>of</strong> <strong>the</strong> size-based inhibition <strong>of</strong><br />

growth: a lower pH possibly lowers <strong>the</strong> speed <strong>of</strong> skeletogenesis and,<br />

consequently, <strong>the</strong> growth rate (<strong>the</strong> soma <strong>of</strong> <strong>sea</strong> <strong>urchin</strong>s is made <strong>of</strong> ca. 90%<br />

<strong>of</strong> mineralized tissues). No matter what <strong>the</strong> cause <strong>of</strong> <strong>the</strong> inhibition may be,<br />

in such simple experimental conditions, <strong>the</strong> relationship between l and τ<br />

appears amazingly simple. A precise measure <strong>of</strong> <strong>the</strong> pH gradient among<br />

aggregates and <strong>the</strong> quantification <strong>of</strong> skeletogenesis speed with pH drop<br />

should bring some more insight into <strong>the</strong>se relations and causalities.<br />

Lacking such information (<strong>the</strong> only experiment on <strong>sea</strong> <strong>urchin</strong>s growth<br />

related to pH was performed on larvae, Bouxin, 1926), eq. 32 is currently<br />

<strong>the</strong> only way to quantify <strong>the</strong> degree <strong>of</strong> inhibition.<br />

Even in <strong>the</strong> present example, eq. 32 seems to be a very simplified<br />

relationship between l and τ. The 10% smallest animals do not follow it.<br />

Pr<strong>of</strong>ile 2 <strong>of</strong> Fig. 34B at 600 days shows that <strong>the</strong> <strong>model</strong> overestimates <strong>the</strong><br />

smallest fraction (and thus underestimates <strong>the</strong> peak <strong>of</strong> mid-sized animals)<br />

at this age. Adaptation <strong>of</strong> l, or even k2, is perhaps necessary to better fit <strong>the</strong><br />

smallest fraction. Again, <strong>the</strong> simplest possible <strong>model</strong> was presented but<br />

many variations can be conceived.<br />

Relation between l and τ could be even more complex in o<strong>the</strong>r<br />

circumstances: different rearing methods, large and small animals <strong>of</strong><br />

different ages and/or genetic origins maintained toge<strong>the</strong>r, periodic size<br />

sorting in culture, etc. Pr<strong>of</strong>iles <strong>of</strong> l in function <strong>of</strong> τ must be studied in each<br />

particular case. It is also probably very different in <strong>the</strong> field, or for o<strong>the</strong>r<br />

species. The <strong>model</strong> still needs to be reformulated and tested before being<br />

used with field-collected data (mainly cohort separation, or mark-<br />

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

171

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