In this study, the formation of the adult sea urchin shape is rationalized within the Turing's theory paradigm. The emergence of protrusions from the expanding underlying surface is described through a reaction-diffusion model with Gray-Scott kinetics on a growing oblate spheroid. The case of slow exponential isotropic growth is considered. The model is first studied in terms of the spatially homogenous equilibria and of the bifurcations involved. Turing diffusion-driven instability is shown to occur and the impact of the slow exponential growth on the resulting Turing regions adequately discussed. Numerical investigations validate the theoretical results showing that the combination between an inhibitor and an activator can result in a distribution of spot concentrations that underlies the development of ambulacral tentacles in the sea urchin's adult stage. Our findings pave the way for a model-driven experimentation that could improve the current biological understanding of the gene control networks involved in patterning.
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Figure 1. Case of no growth: $ r = 0 $. Bifurcation diagram in the $ (c_1, v_e) $ plane. The branches of the spatially homogeneous equilibria $ (u_e, v_e) $ for model (16) are shown as functions of the parameter $ c_1 $. The parameter $ c_2 $ is fixed as $ c_2 = 0.04 $. In this specific case, the trivial equilibrium $ P_0 $ is an equilibrium for all the values of $ c_1 $ whereas the two equilibria $ P_1 $ and $ P_2 $ exist only in the range $ 0.01 < c_1 < 0.16 $
Figure 2. Case of growth: $ r \neq 0 $. In the parameter space $ (c_1, r) $, the region below the curve is where existence of the two non trivial equilibria $ P_1^{(r)} $ and $ P_2^{(r)} $ is ensured. The parameter $ c_2 $ is fixed as $ c_2 = 0.04 $. (left) The case $ \omega = 1 $ (right) The case $ \omega = 10 $
Figure 3. Case of growth: $ r \neq 0 $. The branches of the spatially homogeneous equilibria $ (u_e, v_e) $ for model (16) as functions of the growth parameter $ r $. The parameters $ c_1 $ and $ c_2 $ are fixed at: $ c_1 = 0.03 $, $ c_2 = 0.04 $. The trivial equilibrium $ P_0 $ is an equilibrium for all the values of $ r $ whereas the non trivial equilibria $ P_1^{(r)} $ and $ P_2^{(r)} $ exist only within a particular range of the parameter $ r $ and disappear because of a saddle-node bifurcation. (Left) The case $ \omega = 1 $, (right) The case $ \omega = 10 $
Figure 4. Effects of the growth parameter $ r $ on the Turing region of the equilibrium $ P_{2} $. Turing regions in the parameter space $ (c_1, d) $ are shown for different values of the parameter $ r $. The other parameters are fixed as: $ c_2 = 0.04 $ and $ \omega = 1 $. Turing regions are bounded by the solid curve, the dashed vertical line and the dash-dot vertical line. Top row: (left) $ r = 0 $; (right) $ r = 0.001 $. Bottom: $ r = 0.002 $
Figure 5. The stationary case: $ r = 0 $. LSFEM numerical solutions $ v({\bf{x}}, T) $ of the model (16)-(2) on the oblate spheroid with $ f_0 = 1.9645 $ and $ \xi = 4.1134 $ attained at the corresponding integration times $ T = [0, 20, 100, 500, 1500] $. The model parameters are chosen so that $ (c_1, c_2) = (0.03, 0.04) $, $ d = 0.05 $. The size parameter is $ \omega = 10 $
Figure 6. The slow exponential evolving case: $ r = 0.001 $. LSFEM numerical solutions $ v({\bf{x}}, T) $ of the model (16)-(2) on the oblate spheroid with $ f_0 = 1.9645 $ and $ \xi = 4.1134 $ attained at the corresponding integration times $ T = [0, 20, 100, 500, 1500] $. The model parameters are chosen so that $ (c_1, c_2) = (0.03, 0.04) $, $ d = 0.05 $. The size parameter is $ \omega = 10 $
Figure 7. Spatial pattern in the biological plot related to the variable $ v({\bf{x}}, T) $ of model (16)-(2) at $ T = 1500 $. The oblate spheroid parameters and the model parameters are fixed as in Fig. 6. (Left) stationary case, $ r = 0 $. (Right) slow exponential evolving case, $ r = 0.001 $
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Case of no growth:
Case of growth:
Case of growth:
Effects of the growth parameter
The stationary case:
The slow exponential evolving case:
Spatial pattern in the biological plot related to the variable