Bubble packing, eccentricity, and notochord development.

This paper develops a theoretical basis for the observed relationship between cell arrangements in notochords and analog physical models, and the eccentricity of their cross sections. Three models are developed and analyzed, of the mechanics of cell packing in sheaths. The key ratios governing the packing patterns and eccentricity are cells per unit length λ, tension ratio Γ, and eccentricity e. For flexible and semi-flexible sheaths, the optimal packing pattern shifts from "bamboo", with a symmetric cross section, to "staircase", with an eccentric cross section, at a critical value λ = 1.13. In rigid tubes, this threshold is lowered as imposed eccentricity is increased. Patterns can be observed which are not optimal; pattern transitions may occur below or above the critical λ values. The eccentricity of staircase patterns in flexible and semi-flexible tubes is found to be dependent on the tension ratio Γ, increasing as sheath tension decreases relative to interior cell tension. A novel "serpentine" packing pattern appears for low Γ near the critical λ. The developmental utility of enforcing notochord eccentricity is discussed, as well as potential mechanisms for such control.